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Published by ihe Museum Board 
Adelaide 1985-1986 



Nos. 1-5. Published May 1985, 

No. 1. A bibliography of Aboriginal archaeology in 
South Australia. (Philip G, Jones) ......... L 

No. L A new and interesting species of Cerhegus 
Thoretl (Ischnotbehnae, Dipluridae) from South 
Australia. (Robert J. Raven) . . 15 

No. 3. Millipedes from Australia, 5: Australio- 
somatini from South Australia, with a note on Line 
status of Polydesmus inrwlatus Karsctu and first 
record of a second M ed i t erranea n j ul i d in 
Australia (Diplopoda: Polydesmida, Paradoxo- 
somatidae & Julida, Julidae), (G\ A- W, Jeekel) 

No. 4. Sarcoptiformes (Acari) of South Australian 
soils, 4. Primitive oribate mites (Cryptostigmata) 
with an extensive, unfissured hysteronotal shield 
and aptychoid. (David C, Ue) . . 39 

No. 5. Mollusc t y pe- s pec i men s in t he So ut h 
Australian Museum, 2, Gastropoda: Conidae, 
(Wolfgang Zeidler} 69 

Nos. 6-10, Published June 1986, 

No. 6. A revision of the Irnglophilie genus Brises 
Pascoe w i th a d iscu ss ion of I he Cy phalein i 
(Coleoptera, IfenebrionidaeX (E. G. Matthews) 

.. , 77 

No. 7. Additions to the collembolan fauna of Heand 
Island. (Penelope Greenslade) 91 

No, 8, Mollusc type-specimens in the South Aus- 
tralian Museum. 3. Polyplacophora, (W, Zeidler 
and K. I.. Gowlett) , 97 

No. 9. A revision of the southern Australian starfish 
genus Neciria (Asteroidea: OreasteridaeJ, with the 
description of a new .species. (W. Zeidler and F. W. 
Rowe) 117 

No. 10. Redescription of the mysid crustacean, 

Notomysis austmliensis (Tattersall) Comb. Nov., 

representative of a new genus. (Karl J. Witttnann) 


Nos. 11-14. Published May t98d 

No. II, On TrombeHa aipha n,sp, (Acarina; Trom- 
bellidae) from Australia: correlation, description, 
developmental abnormalities, systematics and 
possible auditory structures. (R. V. Southcou). 

No, 12, The genus Odontacffrus (Acarina: Trom- 
bkulidae). IL Observations on the life history and 
morphology of Qdoniacarus swam n,sp T and 
related forms (R. V. Soutbcotl) .......... 169 

No, 13. History of the discovery of Speieognathus 
austr&tis Womersley (Acarina: Trorn bids formes), 
with notes on its natural history and behaviour, 
(R. v. southcott) - 201 

No. 14. Description of Qdontaamts veifchi sp. nov. 
(Acarina: Tronibiculidae), (R, V r Souihcolt) 213 

No, 15. Published August 1986. 
A checklist of helminths from Australian birds 
(Patricia M. Mawson i I.. Madeline Angel and S. J. 
Edmonds) , 219 

Nos. 16-19. Published November I9B& 

No, 16, Anatomical notes on the land snail ISoth- 
rietfibryon (Fulmonata: Bulimulidae) from South 
Australia and Western Australia. (Ron C\ Kershaw) 

, . , . t . 327 

No. 17, The Epherneroptcra (mayflies) of South 
Australia, (P. J. Sulcr) 339 

No, 18, First representative of the order Macro- 
stomida in Australia (Platyhelminthes,, Macro- 
stomidae), (Ronald Sluys) . 399 

No, 19. A reassessment of the Papuan subfamily 
Asterophyryinae (An una: Micro hylidae). (Thomas 
C Burton) 405 


See Patricia M. Mawson 219 

A rcassessmenl of the Papuan subfamily Astero- 
phyryinae (Anura: Microhylidae) 405 

See Patricia M. Mawson . . . , , 219 

See Wolfgang Zetdlef . * * 97 


Additions to the collembolan fauna of Heard Island 
. 91 

Millipedes from Australia, 5: Australiosomatini from 
South Australia, with a note on the status of Pofy- 
desmus umot&tus Karsch* and first record of a 
-second Mediterranean jutid m Australia (Diplopoda: 
Polvdesmida, Paradoxoaomatidae & Julida, Julidac) 
.,,"., . 19 

A bibliography of Aboriginal archaeology in South 
Australia P 1 

Analomica] notes on the land snail Bolhn'embryon 
(Pulnionata: Butimulid&e) from South Australia and 
Western Australia . . 327 

Sareopti formes (Acari) of South Australian soils, 4. 
Primitive oribate mites (Crypiostigmata) with an 
extensive, un fissured hysteronotal shield and 
apiyehoid 39 

A revision of the troglophilic genus Brises Pa&eoe 
with a discussion of the Cypbaleini (Coleoptera, 
Tenebrionidae) 77 


and S. J. EDMONDS 
A checklist of helminths from Australian birds 219 

A new and interesting species of Cetftegus Thorell 
(Ischnoihclinae, Dipluridae) from South Australia 
.., 15 

R0WE, F. W, 
See Wolfgang Zeidler 117 


First representative of the order Macros mini da in 
Ausiralia (Platyhelminthes, Macrostomidae) . 399 


On Trombelia alpha n.sp. (Acari na: Trombellidae} 
from Australia: correlation, description, develop- 
mental ab norma lit ies s sy stemat ics a nd po ssib le 

auditory structures , , P 145 

The genus Odontacaws (Acarina: Trombieulidae), 
II, Observations on the life history and morphology 
of Qdotttacanw swam n.sp. and related forms . . 169 
History of the discovery of Speleogmttkus australis 
Womersltry (Acarina: Trombidi formes) , with notes 
on its natural history and behaviour .....,, 201 
De sc ri pt i on o ( Ode n mcants veitch i s p. no v. 
(Acarina: Trombiculidae). . . , 213 

Tbe Ephemeraptera (mayflies) of South Australia 


Redescription of the mysid crustacean, N&tomysis 
ausirafiensis iTaiXtra&M) Comb, Nov,, representative 
of a new genus 1 39 

Mollusc type-specimens in the South Australian 
Museum. 2. Gastropoda: Conidae 69 

Mollusc type-specimens in the South Australian 
Museum, 3 r Polyplacophora „ 97 


A revision of the southern Australian starfish genus 
Neetria (Asteroidea: Oreasteridae), with the 
description of a new species . ... . 117 



Aboriginal , 1 

Acari 39 

Aearina 145, 161, 165, 213 

alpha, Trombella 145 

Anura . , , - 405 

Archaeology I 

Asteroidea , , , . . 117 

Aiterophrvinae , 405 

australiensis, Notomysis 139 

AustraliosomaHnae . . , , , 19 

austraiis, Speleognathus , r , . . 201 

Birds (helminths from) 






. 77 


Cethegus , , 15 

Coleoptera . , ■ , . . . . 77 

Colkmbola (of Heard Island) 91 

Conidae . . 69 

Crustacea . , , „ 139 

Cryptostigmata , , + * . 39 

Cyphaleiiii . 77 



Gastropoda . . . 

. 19 

. 15 

. 69 

Heard Island (Collembola of) 91 

Helminths . . , , . . . 219 

innotatus, Polydesmus 
bchnotbelinae ...,,.. 


Julida 19 

JuJidae . 19 

Land snail 


Macrostomida , ,•♦♦...*« 399 

MacrMlomidae ► .** 399 

Mayflies (of South Australia) B 339 

Microhylidae , 405 

Millipedes (from South Australia) 19 

Mites 39 

Mollusc (type-specimens) 69, 97 

Myfcid r .. , 139 

Nectria . . , 

Odontacarus . 

Greasteridac . . 
Oribate mites 






Paradoxosomatidae 19 

Plat y helm iuihes ..,,.,, 399 

Polydesmida . 19 

Poiydesmu* 19 

Polyplacophora . , , . 97 

Pulmonata 327 

Sarcoptiformes . , 39 

Speleognathus 201 

swani, Odontacarus . 165 

Ifenebrionidae 77 

Trombella 145 

Trombdlidac , . 145 

Trombiculidae 165, 213 

Trom bid i formes 201 

veicchi, Odontacarus 


RECORDS of the 



MAY 1985 







by C. A. W. JEEKEL 





North Terrace, Adelaide 
South Australia 5000 



by Philip G. Jones 


A bibliography of South Australian Aboriginal archaeology, indexed accordingly to subject and 
region, and citing 600 references, including published and unpublished material is presented. 


Compiled by 

South Australian Museum, North Terrace, Adelaide, South Australia 5(KK1 
(Manuscript accepted 30 August 1984) 


ioni-S, R G. 1983, A liibliography of Aboriginal archaeology in South 

Australia. Utv. S. M<$ Mb* V)\\) 1-14. 

A bibliography of South Australian Aboriginal 
archaeology, indexed according to subject and region, 
and citing 600 references, including published and 
unpublished material is presented. 











Origins and General rods 



Trends and Method* 


SUBJECT iisi INii 



Incised Stones, CylCOtM 



Stone Tools and Industries 



Stone Arrangements, Structuics 



Rock EngfftViDgJ and Paintings 



Caves and Rockshcltei.s 



Shell Middens 



Bone Points, Wooden ArUfflCtfl 



Canoe 1 recs. Scarred trees 



Excavations of Rinnan Remains 


(10) Radiocarbon Dales 




1 iL'iiei .tl 



Western, North-Western 



Eastern. North Eastern 



Mid-North, Minders Ranges 



Lyre Peninsula, Yorke Petm >i > 



Lower North, Adelaide Region, Fleuricu Peninsula 



River Miliary Region 



Lakes, Coorong. South Lasl 



K. in l'. mx> Island 



Archaeological knowledge is constructed from the 
available evidence; this bibliography represents an 
attempt to document this evidence as it relates to South 
Australia and to make it accessible to researchers. 

The bibliography contains a full listing ol references 
relating lo the theory and practice of Aboriginal 
archaeology in South Australia. It cites published and 
unpublished material extracted from a wide range of 
sources. Unpublished material has been chosen 
selectively; only notes or documents of direct 

significance have been included. Recent newspaper and 
magazine articles about archaeology arc numerous and 
vary in quality. Only significant examples have been 

The bibliography falls into three main sections: 

A. General 

Section A is an introductory section covering relevant 
legislation and general texts relating to theory, method 
and developments in South Australian Aboriginal 

B. Subject Listing 

Section B contains subject-related references 0*Roek 
Engravings and Paintings", "Shell Middens" etc.) 
organized under ten subject headings. Individual 
entries are cross-referenced to the regional divisions 
dealt with in Section C by means of alphabetical key 
letters which follow each entry. 

C. Regional Listing (sec Fig. I) 

Section C includes references which relate to 
particular areas in South Australia rather than to 
specific archaeological subjects as dealt with in Section 
B. These references have been arranged under a general 
heading, relating to the State as a whole, and under 
eight regional headings (a-h). These broad regional 
divisions reflect a convenient organization of the 
reference material. They do not represent precise 
geographic or topographic boundaries or cultural areas. 

The subject and regional listings have been used as 
the most effective means of arranging the quantity and 
diversity of reference material. The two listings are not 
mutually exclusive; regional listings in Section C should 
be supported by relevant material from Section B. 

For those conducting archaeological research in 
specific localities there arc two additional sources which 
may be consulted. Both contain restricted information; 

(1) The annual Bibliography ot Consulting Reports 
prepared by the Australian Association of Consulting 
Archaeologists Incorporated. This can be obtained 
through the Association, P.O. Box 214, Holme 
Building, University of Sydney, N.S.W. 2006. 

(2) South Australian Archaeological Site Reports. 
Two different files of these Reports are maintained, one 
in (he South Australian Museum (Anthropology and 
Archaeology Section) and the other in the South 
Australian Department of Environment and Planning 
(Heritage Conservation Branch). 

The writer wishes to thank Mrs Joan Murphy for 
her cooperation and patience in typing this manuscript. 

REC. S. M SI Ml S 1') ll): I H 

\1a\\ /fl$J 




Am. Antiquity 

American Philosoph. Soc. Mem. 

Annals Aust. Coll. of Dental Surgeon-. 

Ann. Rev. Amhrop. 

Archacol. in Oceania 

Arch. Phys. Amhrop. in Oceania 


Asian Persp. 

Aust. Ab. Stud 

Aust. ArchaeoL 

Aust. Archaeol. Assoc. Newsletter 

Aust. J. of Botany 

Aust. J. Sci. 

Aust. Mus. Mag 

Aust. Mus. Mem. 

Aust. Nat. Hist. 

Aust. N.Z. Ass. Adv Sci. 

Aust. Qual. Newsl. 

Cam. U Press 
Curr. Anthrop. 



J. Arch. Science 

J. Anthrop. Soc. S. Aust. 

J. Biogcogr. 

J. Roy. Anthrop. InsL 

J. Roy. Soc. W. Aust. 

J. Soc. Oeeanist 

Nat. Hist. 

Nat. Mus. Vic. Mem. 

New Quart. Cave 


Proc. Prehistoric Society 

Proc. Roy Geog. Soc. Aust., S. Aust. Branch 

Proc. Rov. Soc-. Tas. 

Proc. Roy. Soc. Vic. 

Rec. Aust. Mus. 

Rec. S. Aust. Mus, 

Report Aust. Assoc. Adv. Sci. 

Rep Wash. GeoL Cong. 

Revue Ethnogr. Tradit. Pop. 

Site Rec. Newsletter 

S.A. Mus. A. D. 

S, Aust. Nat. 

S. Aust Ornithologist 

Syd. Univ. Spel. Soc. J. 

Trans. Br. Cave, Res, Ass. 
trans. Roy. Soc. S. Aust 

Vic Nat. 

Aboriginal and Historic Rcltcs Uuti relocated from South Australian Museum in 1979 
Currently operates as pari of the Heritage Conservation Branch, Department of 
Environment and Planning, South Australia 

Australian Institute of Aboriginal Studies Canberra 

American Antiquity 

American Philosophical Society Memoir 

Annals of the Australian College of Dental Surgeons 

Annual Review of Anthropology 

Archaeology in Oceania 

Archaeology and Physical Anthropology in Oceania 

The Artefact 

Asian Perspectives 

Australian Aboriginal Studies 

Australian Archaeology 

Australian Areheological Socicts Newsletter 

Australian Journal of Bounv 

Australian Journal ol Science 

Australian Museum Maya/me 

Australian Museum Memoirs 

Australian Natural History 

Australian and Mew Zealand Association lor the Advancement of Science 

Australian Quaternary Newsletter 

Cambridge University Press 
Current Anthropology 

Environmental Impact Statement 

\ riends of the South Australian Museum Newsletter 

Journal ol Archaeological Science 

Journal of the Anthropological Society ol South Ausnaha 

JOUfnal ot Htogeography 

Journal of the Royal Anthropological Institute 

Journal of the Royal Society of Western Australia 

Journal de la Socio c Oceanistes 

Natural History 

National Museum or Victoria Memoirs 

New Quarterly Cave Research 

Ovfonl Umvetshv Press 

Proceedings of the Prehistoric Society 

Proceedings of the Royal Geographical Society oi Australia, South Australian Branch 

Proceedings ol the Royal Society of Tasmania 

Proceedings of ihc Royal Society of Victoria 

Records of the Australian Museum 

Records of the South Australian Museum 

Report ot the Australian Association for the Advancement ol Science 

Report ol the Washington Geological Congress 

Revue D'Ethnographie el des Traditions Populaiis 

Site Recording Newsletter, Heritage Conservation Branch, Department of Environment 

and Planning, South Australia 
South Australian Museum Archival Documentation 
South Australian Naturalist 
South Australian Ornithologist 
Sydney University Speleological Society Journal 

Transactions of the British Cave Research Association 
Transactions of the Royal Society oi South Australia 

Victorian Naturalist 


(1) Legislation 

ELLIS, R. W. 1972. South Australia: Existing legislation and its 
implementation. In The Preservation of Australia's Aboriginal 
Heritage. Report of National Seminar on "Aboriginal Antiquities 
in Australia" 53rd and 24th May 1972, A.I.A.S., Canherra 

EDWARDS, R. 1970. Legislation for the protection of Aboriginal 
relics in South Australia, In "Aboriginal Antiquities in Australia: 
Their Nature and Preservation" (F D. McCarthv, ed.), pp. 
159-165. A.I.A.S., Canberra. 

SOUTH AUSTRALIA (STATUTES) 1965. Aboriginal and Historic 
Relics Preservation Act. Govt, Printer, Adelaide 

SOUTH AUSTRALIA (STATUTES) 1979. Aboriginal Heritage Act. 
Govt. Printer, Adelaide. 

SOUTH AUSTRALIA (STATUTES). Aboriginal Heritage Bill (Draft 
form). Act not yet proclaimed (August 1984). 

WARD, G. K. 1983. Archaeology and legislation in Australia. In 
"Australian Field Archaeology. A Guide to techniques!' 
(G. Connah, ed.) pp. 18-42. A.I.A.S., Canberra 

<2) Origins unci General Texts 

BAGLIN, a, and MULLINS, B. 1969 "Aborigines of Australia" 

Horwitz, Sydney. 
BAIL EY. G. N. 1980. Holocene Australia, fn "The Cambridge 

Encyclopedia of Archaeology" (A. Sherratt, ed.), pp. .W-341. 

Cambridge University Press, Cambridge. 
BASEDOW, H. 1925. "The Australian Aboriginal" IVeee & Sons, 

BERNDT, R. M., and C. H. (eds.). 1965. "Aboriginal Man in Australia!' 

Angus & Robertson, Svdncy. 
BERNDT, K, M n and C. H. J98L "The World of the Eirst Australians!' 

Lansdowne Press, Sydney. 
BtRDSELL, J. B. 1953. Some environmental and cultural factors 

influencing the structure of Australian Aboriginal populations. 

Th- American Naturalist, 87; 171-207. 
B1RDSELL, J, B. 1957. Some population problem* tmohing 

Pleistocene man. Cold Spring Itarhnt fympOSfO "« Quantitative 

Btotozs, 22: 47-69. 
HIRDSELL, J, B. 1967, Preliminary data on the trihybrid origin ot 

the Australian Aborigines. Arch, Phvs Anthrop. in Oceania, 2: 



BiROShl l, J. B I 1 )"". The icuilibimion of a paradftgiri fbl The tiiM 

peopling of Cirdilcr \uMr.ilia. //* "Sunda and Salmi* prehiMone 
.: i i -t-iintcasi Aaa, Melanesia and Australia:' (I Allen, 

i Ism. and R. Jtotwi Odtj, pp ll?-W7, ftcMttnw PIW. 

BLAINi Y. G N. 1^75 'triumph ol .he Nomad* A History of 

Ancient Australia!' Macmillan. Melbourne 
CAMPHtLL, A. H. 1967. Aboriginal traditions and ihc prehistory 

of Australia, Mankind, 6 fit -481 
CASLV, li A. 1940. The present state of our Knowledge ot ihe 

archaeology 91 AnsifaJia to "PwceodUTgs of The I hud Conyre^s. 

o| i'ivhi-.turtaiv. of the Far LaM". Singapore 
[ OTTON, ft t . led ) IY66. ■•Aboriginal Man in South and t eniral 

Au-.iialia!' Pari 1. Govt Pooler. Vlclauli 
1 orris, P J. I, I9Q&. "Icatures of prehistoric campsite in Australia'.' 

\hnKoul. ft; 33K-346. 
DIAMOND. J. M. 1^77 Dismbuiional strategies. /)/ "Sunda and 

Sflhtil- prehistoric stUiiies itl Southeast Asia, Melanesia and 

Auslrahu" (J. Allen. I L.uUoit, ..mi k Joik-s, eds.|, pp. 1$5 310. 

Academic Press, London, 
LDWARDS, R .(ed.L 1975 "The Preservation ol Australia': .Aboriginal 

Heritage. Report of National Semtnap an ftbttrigthal Am in; nines 

in Australia, May \&T£? A.I.A.S., V anhena. 

FLOOD, I P>83. "Arvri.M;-oh» H s nlihc OiuHii v.--, i.ollins, Sydney. 

• ill I, L L). IW3, Ausiralian Aborigines and {he giant o&UnCl 

marsupial*. *1/W- Nan Wto 14 263-266. 
OOULD. R. A. 1973 An&rratiini archaeology in ecological and 

ethnographic perspective Mister Modular Publication.!: 1-33. 
QR1 IV WAV I. I%2- AahjLu'MO-". m Australia. Southwestern tore, 

IH: 25.10. 
HAMMOND, (.. I^HI, Aspects o) leeai Mer<ii'ie.UK>-- in Aichaeol' ' 

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MORTON, i) ft. 1981. Water and woodland: the pcopluie of uIili 

A I. W \rw\tflfc1 |i 21-27. 
HORTON, D. R. I98L Lad) thought on early m..r. |M Aum I I 
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AjSTOM, O iwv Magic -tones oLthe irihpf SKI and norirwa-,i ftf 
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HI A( K. M I 1942 "( v!,ons. the Mv, ( rrv Ston^ ol ihe I.-..H ( 

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\V/ ( . S AUM- M; ll-lft, b 
i 'AMPHPI L , I- P 1924. An HCCQUI1I ut H liiihciui iiiircciHdctt (yftc 

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i. CM 'l*Li%. 1 1. M. I'M?. Incised stones ol South Australia. Mankind. 

M 2V2 29K. c 

r Itukl I.. I. ||i i- . -h n 1.'i D i m riv-ir- lUtfid In V- itint L 

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MOliN I t-OKD, C. P. 1938. Engraved pebbles* from South Australia 

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MOUniiokd, c. p 1940. An Aboriginal engraved pi-hblc from 

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MOUNTFORD, C P. I960. Unusual nn.t. mfkim '" Australia. 

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PREISS. K. \. I9fi3, An interesting irane artefaci nom the Pllflden 

K.mvrs, V \itsi. Nut M S3-57, C 
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phi 1 1 ime k tt. i92»». Cylindnvcpnioal woncs frron A»eonu:.. 
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AJSTON, G. Letters in W. H. Gill 19201940 3v. Unouhl^hcd Ms 

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ALICK1N. U. W57. Ausiialian stone industries. r>ast and present ./. 

A'ov. Anthrup. fast H7 USHr". a t I, « 

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CLAKkF, C-. ,1. 1979 An imparl bud IT OH a Bondi point horn Koodf 

Drift, Souih Ausiialia AneiiKt 4: <*i8-72. g 
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Ai.isli.llu. S Aust \af 25 FH. C, fe, n 

t'OOPliK, H. M- |95(J Stone iniplctnenli from a mangrove swamp, 
So.nh CHtnelB Ttanx Rtn. Sat & 4/ysi. 73. 220-222 c 

I OOPFK, H. M. 19S.F Maieriuleulnucot ihe Australian Aboriginals. 
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\tu\ H: ^Fioi. b, e 

COOPl K, IF M 19^-9 I jrncdttltjcoloyicjil sl.if.e implcntepls Iroui 

HallutCove, StH.ili *\iiv.triii.;. ]ran\ Rum SMk £ .-1^/. «2c 55-60. r 
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AV', K -bys/. Mits. 14: | -IS. \ 
•: ' h )Ptk, if M. 1961. Archaeological stone mqitcmeMH alony ihe 

Lower River Wakebeld, Souih Austmha fntn\ Kav. Sot: S. AtlsL 

84. 105-115 b 
CXXWHHi H. M I96r. Arehaeolopieal stone futplemcntj from a 

tagjDun heil. Kangaroo FJand. South Australia. «cC 5. .-to. \/us., 

t5. 109-327. h 
t.Ot )PLR, IF M 1%K A tuitliui desenptum .a "Imi^- stone implemenis 

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EDWARDS, K I96C PieiiiiiHUuy survey of I he Anonym;. I remfonn 

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EDWARDS, R. |y64. Rock engnwrngi and stone ifiiplcnienis of 

Pilcairn Station, Nonh-ea-acrn South Australia, to; L Aim. Mivl 

14: M344I. t> 
LDWAKOS. K 196f» Ausluhfes used tt>r Aboriginal iniplemeiiK .11 

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E1ENNIKEN, I. J. ^nd WHITE. I, P 198? Heat treatment ot 

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CiOUl U R. A. 1966. Some Mont artifacts ol rhc Wonkanguru ul 

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GOUED, K. A., KOS1LK, f>. A., and SONTZ, A H. I. 1971. The 

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4m. \ttti<tutty 36: 149-169. a 
HAKVt ; Y, A- I94C Klitu implements of Tasniunian niamilucunr found 

ai Cape Hart, kat^an.o l.-.taud far £ - -lust. \!ns. f»: <oC3f«N. h 
HAY DUN, B- D 1976. Australian Western Desert lithie teehiinlogy; 

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Pl.,0. thesis, Lhiiv. of Toronto, a 
HAYDEN. b\ D. 1977, Stone tool tUntiHODS II lilC Wr,i«.rn iX-sen 

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HAYDEN. B D t9SE Paiacolilhiv. Reflection,. | i|foi feci |0& ,-md 

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HEWITT, R. 1969 and 1971. Analysis of implement*, from Eucolo 

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HEW17 r, R. I97,v The Areootta Plateau an outline o\' Aboriginal 

lubiln's and alio .;. Afifhiop. Soe S. AtiSt. W 6-19. b 
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HBCOCK, P. and HUGHES, R, J l9S3.0nt method pt nMpnliiifi 

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S7 9K. x 
HOWCHIN, W. Igg?, Hie Mane impIe-i'MU oj |||| Adeiaidv ",he 

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HOWCHIN, W ( . 1921. On liic occurrence of Aborigirtftl ><\*w 

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HOWC HiN, W. 1934. "fhcMorie ImplemcniN of H.c AddftOK 1Mb? 

Of Aborigines now extinct" Gillmiihain, Adelaide, t 
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study of eolirlt*; some observations on the natural loiec^ ;ti \Natk 

in Hie production ot Hv»kcd stones on the Leniril AUNiralian 

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I! HINSON, J. i^ I95S. ] -utr lo S. R. Mitchell re artelacis town \(j 

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KAVIMINGA, t_ 1971 Microscopic and experimental studies ol 
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ihc«?i Univ of Sydney ■* 

KAVIMINGA, J. 19SU A |T)OCUOJ!0&I "ivcMnMhon ol Watatian 
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KAMMIM.A. I IVSI "<jwi iti, i .1 , i miMnal uiaiv i ■ d 

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I .Wll'i.KT, R. 1 1975. A prclinmiaiv report of b0IK< waiMed hhules 
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I AMPtKI. R, T i97^, Vuriahon ... At.Mn.l.H'-^.ie a,>..e 
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I J Mil t Pie and Proiiihisid.iL Symposium, Nice, Sep). 1970. 

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McBRYDE, I. 1977. Prehi-aorv IJlJjJc evidenet. |0t evploilalion a) moiic 
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McBRVDE. I. 1976 Wtl tm ee Moor ring: Or where do ixes come 
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McBKVDL. I., and WATCHMAN \ 1976. rhcdiMribunou ot | ■,. 

stone .iv:'. in gouUlHrasitcm Australia: ■ prelttmnarv rvpdll 

Mankind Itt: If. VI 74. ft 
McCARl H\, I. |> I94E The sione axes of AlistraJJa. AMI. A/u-V. Md$. 

7, p P 223 ■::«. x 
McCarthy, f. u t94i clipped stone irnpi^mcpisol ift< Abonooto 

\UXt \Jus \[uk.. 7; J^-IW u 
MU \RTft>. K U, BKAMLLL, L.. and NOONE, H, V V. p<4(> 

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MCCARTHY, h 1>- 19fi? The tboriglnal past: archaeological jnd 

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McCARTHY 1- D. 1977 The use of stone tools 10 map patterns of 

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McCOURI, I 1975 "Al-torieiiml Anefaw*! " kwmv. \delaide. g 
MAMHhWS. J. M. 1966 Tlu Hoabhinhian athnitie>: Dl 

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HAl Ul*, I Mc<iO\V\S K I Ulld (.11 l| I KM : 's- f I I- WSl shor.gmal 
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VIOl ;N I FORD. L . P. 1426 l%l. Arranged Stones, Sione Stntcn.M.-s, 
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MttUNfLORn, C P. -"id HOWAROS, R, 1062, Aboriginal rock 

engravings ol extinct ereaiui'iS ui Souih Ausiralia. Man fi2: 

9 -U4. u 

MOCJNJKORI^ L. P.. and HOWARDS. R. I%.V Rock engttvjfltf 

ol' Panaratuitcc Staiion. not tlt-ea^iern South AUbtralia fio/i\ Km, 
W., v /(Wff «6: 131-146 I: 
M<U vtlORO, c. P.. and EDWARDS, R. 194*. Rock eugraiinys 
in the' Red GorgCi Lleception Cieck, norlhtrn South Australia. 

Ni >RHS. M. K i'>7.s, An inventory ol Aboriginal shelter painting 

on^o Historic Reserve, South Australia, luly 1977_ /. 
■\rithrufi S<h\ S ,4(/\/. 16: I -My b 

NOfittS. M. F, 1961- Mannahill engraving site (Ratolta). Report 10 
the ,VI AS l,|<esctipi ipoicessedi. A.l \S.. (. anbcua, b 

Ni >BKS. m. \|_ iys2 Aboriginal rock an in Olary Province, SA A 
n-pon on a site recording programme carried out on toimecn 
Abongma! roMi an ftef on die sheep stations knossu a : - 
Rimbowrie. Outalpa, Karolta and Oulmna. Typescript (pnxessed}, 
A.I.A.S. Canbetra. h 

NOliBS, M- l u «4. Roekar. in Olarv fVov. ner. Somh Ausimha. WRA 

tax. £Mii n 

PRLtSS. K. A. 1961 nxk painting', on the South R6t# 

-:.o,e«. S Alt$l. V//f. J5:" J7-8. d 
PRLISS, K, A. I96Zi Aboriginal rock carving. A posMPIe occiurence 

at Point lunon Voike Peninsulj, Sumli Austntha. A Ai/\f \'of. 

Mt_ 42. d 
l*R] JSS, K. A. lv)n2. Aboriginal rock carvings and paintings iir the 

"Canvon" Hinders Ranges, South Austrdhu. S Aim. ~>\ut 1Z) 

2.i -27. I 
PKl ISS, K. A. WM. Aboriginal ro-.l palnlinfi n the lowei MoifW 

l.oltv Ranges, South Austmlia lour more siles ilesenbed. .K 4/a7 

Vfff 39 vl 2. i 
H M-iPL. C. L 1977 koonalda C avc— the beginning ol jriistic 

c\piesMiin. Vrn 1 Outnt- Q«f 2 226-234, a 
SHARPt.CJ ,andSHARPf,K | I ?Tft A preliminary survev of 

cnKiHved boulders ti' ihe an sanciuaty oi Koonalda Caw, South 

Australia. Mankind 10: pp. tjvun, a 
SHLARD, H L, 1927 Aboiiginal rock slteheis and earvuigs-ilinv 

loealitic"- on the l.ouci Mnnav. Trutts. Roy. Sot, .V Aii\t 5\ 

137,140. r 
SHl-ARI), HI. |o27 Ahoriginn! rock carvings A) IJevon Down;,. 

Rbci Munav, South \ustfalia l)ar>\ Rt<\ Sou S Ausi SI: 

IS v). i 

SHI- VRO, Ml, IV2K. Abongiual rock paintings SCTCI1 miles uunlt 
ol' Hlanchetown. Rjvei Vluiu\, South Australia fhmy f<,i\ W 
S 4iff/. 32; 231-234 f 

SlAin tTOI*, P. S 1931, Aboriginal retfo in the Ulinnun Dn.,.u 
lock cai\uigs tandi Aboriginal stone sirnciun-s \ 4USC N&t 

12; 21-24. i 
StIRI INC.. B V- I002- Aborinin,-,! rock pairiling^ou 'In .vmuh K..:- 

l-:r.-r ftamssa Ranges. Trims. Roy: Soc S, \M 26. 20S-2II *• 

ftf* S MJeH Mi is no > iu 

,U/i-. /VAS 

TEUSNER, R- E. 15(63. Ahoi initial C3V? DMJHtflfl ON f|K W«CI v; 

UNI RJen Valley, Soulh Australia. Mankind it: IS 19, e 
TINDAI E> N. B 1935, Rock marking* irt Souih Australia. Aftitauin 

9: SMI c 
TINDaLE, N. B. F951. Comments on supposed representation-. '•' 

giant bin! tracks ai Pimba. tec i .4/vay. \fus. ft 38I-3S2. a 
TINDAI F. N. B.^odMOUNTKORU. C P. 19?*. rv.n,r <-,.,» kinfls 

on rocki ai Morowtc, South Australia i'ran\ Kty Sffc" V fytit, 

50; 156-159. c 
TINDAI E, N. B. and SHfcARD, H. L 1927 ,„a 

paintings. South Para Kher, South Awsiralm. fetfM ftgrji \or V 

AUtf. 51; 14-17- c 
l JCKO, P 1967. Australian rock an in world amicx! A/.d S Nw$t&t& 

2: 44-52 X 
VERBRbGOE, A. R 1970. "Corpus of the Hand Figuration* in 

Primitive Australia" Editions Ophrvs. Cornpieync 
WHJTE, S. A_ (912. Name an and rock slickers In "5. A White. 

Ooldea and other sketches!' Adelaide, 67-74. e 

WRIGHT. H I. MT7& Thcart Of 'he o K.-k-t-i .yruvers /« "Thi Ai.m.-.Iu.- 

Aboriginal Heritage: An Introduction through the *\rts" tR. VI 

Benidt and t. S. Phillips, eds), pp, 128-154 L're Smith, Sydney, x 

(5) Caves and Roikshdfvrs 

\si |N, C P "id ttLDNARIk, H G J9S4. karake Cave— a. 

preliminarv repot i. Artejurt 8: Jj? -3> g 
VSl IN.G 13. and HEDNARIk. R. G. |9feC karlic-rrjoiiipooi Cave 

\ preliminary tcpon /6?eA' Art tesrunh I; 36 -45. 
\si IN, G. U and BEDNARIK. R. O. |9fl4. Koonne Cave, South 

\i.iralia R»>ck All Research 1. 142 -144. a 
BROWN. R. ISfiJp, Koonalda ;cen through the eves of a field-worker. 

.Annuel I Ih 6 7 a 
i OIL J. 1957. Never-never underworld; eights eenfunes AfiO, 

Ahiirii-ine, knew a w.iv 10 a mysterious world below (he Nullarbor. 
miration PQsf, Feb 14, 1857, 10-11, * 
I) '-A II :- V (..and i RANK, R. k 1984. A review ol' Aboriginal and 

European cultural sites relating u> C$*tt and oiflCJ kJ'-t knifes 

on the Nullarbor Plain in South Australia. Report to the Heritage 

Conservation lira neb. SOlirll Australian Dept. of Ln-.uoriniua 

and Planning. Applied Vauiral Rcsoutcc Mnnae^un-m. Canberra 

DUNKLDY. f, R, m VVtt.I i'V l M_ I. (eds.) 1%7. U.jv 

rhc Nullarbor: A review of speleological iives|i)iaHon\ in \h$ 

Nti Harbor Plain. Soinhcin \uMralia,' Univ. '--i Sfpl Sydnw * 
UMAKHS, R. 1969 The treeless plaffli HemtsohM 13; Z4. a 
flAVAROS, K 1V<>V. PrdfiftQrtc art in KoonaMa < ave (Atisoaha), 

BoU. Dvl Ccnlm Cuttntno Oi pnfflt, 4: 117-130.. a 
LDWAKOS, K., and MAYNARft L. I%7, Prelnsto.-e -trf tn Koonaldr. 

Cave. Pfoc. Hoy. Crux Soe. Aust., S, AytfL Krunch 68; 11-17, a 
t:L KIN, A. P. 1971 Archaeoh.igv of Konnalda Caves jl reviev, arnJr. 

Aivh. /'/nv Atuhrop. m Oceania 5: 162-6. a 
| : RAMv, K. (9^1 I he sedimcra-. In "Arebaeologv oi die QuIllU tJiltt 

Koontitda Cave" tR. V S Wn^ht, ed.) ( pp. 3144. A.I.A.S., 

Canberra, m 
IK \NK, Hi IU?I. r .tve icdimerrl.s aspateotnvirinimenial indicalois, 

ttndthl ■ in i Biat> sequence iii KooiuldaCa\e. tn "WlPriginal 

Man and Lrivironmcru in AusiraLa" (D. J. Mnlvaney aJffd J 

Cols.-n, tf4ff T J, pp. 94-104 ANU PfE», Canberra, a 
viAl I 1,S. A, m% Results Ot the I960 Koo.uM.. I ..., |S,v. ( 

L'spediriLMi ofthe AiLhaeoloeieal Socieiv ot Victoria AtVtfiffl 10; 

J -5. a 
ClAtLUSi, A. L%S EwSVflltOns ai KooiraWfl Cave, Souih Australia. 

Curr, imfovn 9: 324-32$ a 
CiALUJS A. 1%^. Pfn.erai art m koor.nld:. Cave, NullarOor Plain, 

S0Uth \u-ir:tha Ih'Hchti' <*: 41-4V- a 
Ct At US. A. 1*MK. AichaeoloL-K.iil cM.iivfiUon, ai Koonalda, Null^rhor 

r'lain, 1957 l%7. 7. Anihrvp. Sor S Ausi <> as u 
CAI 1 1 >S, A l l >7l ( Results "I ilu exploration o* KodTiidda C;ivc 

1056-1968 )n "ArJiaeolovv or the Gallu-^iiu Kooualda C avc" 

(R. V. S. WViehi. eel. I pp 87-1 33 A I.A.S. Canhorr.i. a 
i.AMvilMi, S. l9dS Udderwnrld of ihc Nullarbor. Wtttktthnut .IP 

V0-J9 M 

QR1MWADET, K. I^H. f-\pcdiiion (M.i.niwidiine Cawj. H ild 1 tfc 
l(h 72-77. 90 a 

Hit I . A. I . IBffT- ChcckltM of caves and icla'ed le-nnri-- M "I ..u- 
Of the Null.ii-hor: A lestcv. o1 ipcleolo^ical invc'-luTiMon< in ilur 
NniLvl'or plain. Southern Australia" (J K. DmiMey and T. M. t 
\\\A<i\, ed,U, PP- 50-tiI. Univ of Sydney Sydile-y. a 

HOPi,, I >»., I.AMPtRT, K. I. (OM'^JHSUN, R.. SMITH. M. 

J., ftoO va*i n : TS, G, I- 197? Late Pfe^toceete remain'. 

Iroir. Sr'on roiA ^^ICI Kan^aioo t.sland. South Ausiraha ./ 

HtQWW- •» 36J'3B5 h 
IAc KA, H I9?4. lincf [e^tfri on humai' ttfflaViC tf| Woniiuu I 

Nat.ieooite. S.A. Cave t.\piw'Qn\>r, fjfW/3 ''S.V; \m<irtt<r 18, 

JENNINGS, J N. I97y, Man and other an.mals m Au'aralian cave 

and shelters; a review. 7m/i\ //r Doc Wt'.s ^U\ 6; 93-130, a, f, g, h 
I ANR, E Ah Slid RICHARDS, A. M. 1966. Handpamungi in tflvca 

with special reference to Aboriginal hand stencils from caves on 

the Nullarbur Plain, southern Australia. Hflicnte A: H-5U a 
i .AWLLR Dt iMt Wonderland under the Nullarbor NWW £a/e l«- 

336- W?. a 
MARTIN, A. 1973, Palynology 3nd ht^lorieat ecology Of WflM ■ BVC 

CXcavartDns in the Ausirjlian Nullarbor. Ami, J. of Botany l\ 

283-316. a 
MARTIN, S. 19*2. Nullarbor Plain sitex Aho/wno! Uernm* . ./ ■ 

5- f> {S.A. Ocpanmont of Lnvironment antJ Plannint'i a 
MARUN, L. H. 1970. RepoP on Nulbrhor litld trip m f/inuary- 

february, (A.I A.Su Doe 70/919) Processed, a 
MARUN, L H. P/72. The Miniing and their prcdeces.',or.s on the 

coastal Nullarbor Plain. Ph.D. thesis. Umversiiv 0\ Svdnev. a 
MARUN, L H and THOMPSON, P. 1969. Progress report on prTDjcCl li^- (A l-A.S. Ooc 6y'S7«) Processed, a 
MASEY, T. A. 1882. The red ochic caves of the B&U& |W R»t 

Augusta Dispatch and t tinders Advertiser, June 9, 3, cots< a, b e 
MATTHEWS, P (ed.). 1%8 Sj»cleo Handbook. Aust-,n. Spet 

Fcderauon, Svdnev a, C g, h 
MAYNARD, L., and EDWARDS, R 1971. Wall Markings, tn 

"Archaeology of the Ciallus .Site, Koonalda Cave" |R. V. S. WrigltL 

ea\). pp 6l»80, A.l A.S , Ccmben.. a 
MLGAW, J. V. Sv 1%6. Theeihnographie and archaeological eonlept 

ot Koonalda Cave *p (A.l.A S. Doc 66/455) Proce\sed. a 
MULVANL'Y, D J. 195$. Murray rock-shelter's Stone- A<>e seuets 

Ktvcrlantier. Sept tV5H. l.V 29. f 

mulvanuy, d x iy5M. they fived tn the Valley 4,oou jwirt agn 

RiHrtander. AUfi TO8i ^- 7 f 
MUIVANHY. O. J. 1959. What Valley hunter* are 5,000 years aga 

Rnrtiandct, Sept 1959. 8; 3t». f 
MUIVANF.V, D. J. M0*. Archaeological excavarions ai FracORJ - 

l^ndin^onthe Ijowci Murray River, Souih Australia /^/rvt Koi: 

Voe. 1 7c 72: 53-85. f 
MULVANCV, J., LAWTON, Cr. H., and TWIOALE, C R I9f»4. 

Archaeological excavations of rock shelter No. 6. Promm's 

Undine, Soulh Australia. Proc, Roj: Soc. Vic. 71: 479-516. f 
MULVANtV. D. J. 1969. Ancienis of tlic Murray Valley. Rivertander, 

Dec 1069. 6: M.t 
PARKIN, L. W 1938. A limestone cave at Swan Reach. & Aust. Nat. 

19: »V9. r 
PRETTY, G. L. I960. TbcCEjG. <is.A ) Nullarbor expedition 1959/60 

aahaeototfica! notes. Svd. Univ. Spct. Soc J, 6; 2-7. a 
PRETTY, G I i and CALLUS. S. A. 1967. The anthropology And 

archaeology of the Nullarbor Plain, tn "Cave> oi' '.he Nullarbor 

j review of speleological investigations in the Nullarbor Plain, 

South Australia" f J. R. DonMey. =*nd T. M L. Wjgtey, eds.) t 

pp 47 49, Speleological Research Council, Sydney. » 
PRICE, A O. 1952. St Mark's College jcientific work at Fromm's 

l^andine. Pruc Geog. Sot. Aust.. S. Ausl Branch *I3: 25-27. f 
SHAKPL L-- L., and SHARPE, ^. J. iy76 A preliminary sttfW 

engraved boulders tn the an sanetoarv of KOOpaldS Cave, South 

Australia. Mankind 10: 125-130. a 
SHARPf . C ^- 1977 Koorvatda Cave— ihe beginning of artistic 

exprcssionv iwv Quart. Cave 2: ^36-234 a 
SMI f H, M. A. 1977. Devon Downs reconsidered— an otcrcuc in bio- 

archaeotoK.v- B.A. Hons. thesis, ANU, Canberra, t 
SMITH. M A. 1978 A note on the fauna from the Devon Down*. 

Sbeftar Ausi. ArchaeoL B: 19-21. f 
SMITH, M. A. 1982. Devon Downs reconsidered changes Ifl site use 

ar a Lowci Murray Vnlley rockshetter. ArchaeoL tn Oceania 17 

Ji.ry-116. f 
STOCKTON. ) 1*^83. Ob^ervationv on ptchi'.rone and hisione uw 

of Nullarboi C^ves^ Compiled by Jim SiocMon horn observafiOfiS 

by p. Oillieson, S N. Jennings, A. Smile and J. SiocRion during 

a nip to examine sand duner, and Jave 'edimenls, A.H.R, 

(Restrieied». fl 
THOMPSON. P. 1970. Report on fieldwotk on the NulUrtxv pfaf?, 

SA, (A I A.S. Doc 70/972) Processed, a 
THOMSON, I. M. 1947, Nullarbor eaves. Walkabout U: 
THORNE-. A. O. W7E The Tauna fn "Archaeology ofthe Gatlus Site, 

Koonalda Gnff (R. V S. Wright, ed.), pp. 45-47 A.I.A.S., 

Canberra, a 
J tNDAVE, N B 1933 Tanunoola Caves, Situth-easi of South 

Aushalia. e*eoto^ie-at a/Id fthysiogta^htC^I noies. Tmns, Ror. Soc 

S *M 57; 130-142. a 
TINDALE, N. e. and MOUNTFORD, C. P. 3936 Results of The 

excavation of Kongerati Cave mar Second Valley, South Ali$tt$ija, 

tiec- S AU$t. Mtis. 5. 487-502. c 
WCXJLE O. 1940. What the eaves tdl of the Nullarbor Plain. Wttdtuv 

WRICH r, R. V, S (ed). "Archaeology of the Oallus Site. Koonalda 
Caw'.' A.l.A S-i Canberra. » 


<6» Shell Mtdfiem 

AMBROSE, W K, 1967. Archaeology and shell middens Arch Phys. 

Anthrof*. m Oceania 2 169-IK7. g 
BAILt v, u K 1975. The role of molluscs it) constat economies; Hie 

icsull of midden analysis In Australia. J. ofArchacoi Science 2 

45 tiu p. 
BOWMAN. G, and HAKVHY, N. 1983. Radiocarbon dmiflfi marine 

siiells in South Australia. 4WST Archueol. 17. 113-123. x 
E 1 MER1DOE, R Jnr. 1893. The "Minm Yon?" heaps at the North- 

WKi Rend of the River Murray. Trim*. Rov. Sen: S. Aust 17: 

21-24. r 
GILL, L. I> 1974. Aboriginal menus and cooking methods as interred 

I'rom atvhacoioaical sites in south-east Australia. Artefwf 35 1-8. g 
Oil l.LSHll , R., and POT ACH H A. 1970. The suiiabilily of marine 

shells lor radiocarbon dating of Australian |.iehisior>. proceedings 

uj ttn' International Conference on Radiocarhon During, Univ. of 

Calif. July 1976. y 
< ill LKSPIL-, R , and I EMPLE, R. B. 1977. Radiocarbon dating of 

shell middens. Arch, Phys. Antfirop. in Oceania 12: 26-37, g 
UJEflBERSj R- A. 1978. Meals and menus: a study of change in 

[Hrluston. coastal SfttftemertM in South Australia- Ph O I bests. 

ANb, Canberra, g 

(7) Bone Points, Wooden Artefacts 

EDWARDS, R. 1968. An Aboriginal bone sheathed point from 

1 romm's Landing. South Australm. Monknul ft: 090*691 f 
LUEBBERS, R. A. 1975 Ancient boomerangs discovered in South 

Australia. Nature 2S3: 3V. g 
LUEBBLRS, R A. 1977. Progress report on the mending of the Wyrie 

Swamp wooden tool assemblage UflpUO M$. A I A S . 

Canberra g. 
MCCARTHY. fr. D. I940 The bone point known as muduk in eastern 

Australia. Rec. Aust. Mas. 20: 313-319. f 
Met AKTHY, I O- 1952. A Werpoo, or bone dagger, from South 

Australia, AwA Mus. M<& w. 290-292 
MITCHEI L, S. R. 1958. An Aboriginal bone industry, Mankinds: 

194-199. f 
MULVANEY, D. .1. I960. Archaeological excavations at Fannin's 

Landing on the Lower Muxra\ River, South Australia. Proc. Rov, 

$OC Vic. 72; 53-85. f 
TBUSNER, R E l%5- Repnr' ol dKt.tivi.-ry ot a bone-point near 

Morgan. J Anthrvfi. Sen. £ Aust I. 4. r 

(8) Canoe Trees, Scarred Ttves 

BASEDOW, H. 1914. Evidence of bark canoes and food earners on 

the River Murray, South Australia. Man 14: no. 63. f 
BLRNtXT, R M_ 1941 The barK canoe of the Lower River Murray, 

South Australia Mankind 3: 17 28. r 
BLACK. E. C. 1947 1 he canoes and canoe tree* of Australia. Mankind 

y 351-361. f 
BLACK, E. C. 1967. Survey Of Aboriginal canoe trees along the 

Murray River in South Australia; progress report. 1p. (A.LA.S. 

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UOEHM, E. F. 1948. Aborifiinul-marked trees in southern districts. 

.S Aust. Nat. 25: tf_i. f 
EDWARDS, R 1966. Aboriginal canoe trees. J. Anihrofx Soc. $. Aust. 

4 4-5. f 
EDWARDS, R. 1967. A unique canoe lice from Manm*m, South 

Australia. FO.SA.M. Newsletter 4; 3-4. f 
EDWARDS, R 1972. "Aboriginal Bark Canoes ofihe Murray Valley" 

Rjgbv, Adelaide. <" 
EDWARDS, R. 1973. Aboriginal bark canoes, Hemisphere. Jan. 1973, 

9-16. r 

MAI E.H M andTlNDALE.N. a 1925. Observations cm Aborigines 
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l l NDSAY, H A. 1957. Canoe trees of the Murray Valley. Riverlander. 
DDC 1957, 21. t 

(9) Excavations of Human Retnains 

RERNOT. R M , and JOHNSTON, T H, 1942. Death, burial and 
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BLACKWOOD, R and SIMPSON, K N. G. 1973. Atmudes of 
Aboriginal skeletons excavated in the Murray Valley region 
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ftK KtE. K. L. I96&. Aboriginal skeleton Hndings at Nildoma /. 
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B/\SCDOW. H. 1913. Burial cusioms in die Northern Flinders Ranges 

of South Australia Man 26: 49 53. c 
HALE H M. 1926 Some Abonginal graves at Adelaide. Suulh 

Australia. S Aust. Nai 8: U. t 
HAt £. h M . and TINOALE. N. B. 1930 Nole^ on some human 

retnains m the Lower Murray Vklles; South Au-stralta. /fci; S Autt. 

Mas 4 145-218. f 
HOWCH1N, W. 1919. Supplementary notes on the occurrence of 

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PREISS, K 1966. Aboriginal skeleton near Murray Bridge, South 

Australia £ Aust. Nat 41 39. f 
PRL1 TY. G. L. 1967. Rescue excavatiptis ol an Aboriginal grave, 

lailem Bend, South Australia. .S; Aust. Nat. 41: 77^80. i 
f'tiLTTY, G. L. 1968 Excavation Of Aboriginal u.iivfs ji C.iidgealpa, 

South Australia. Rec, 5. Aust. Mux 15. 671-677. e 
f'KOKOPEC, M. 1979. Demographies! and morphological aspects of 

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SMhAKD, H-, MOUNITORD, C P., and HACkLTi, [_ . j. IV27. 

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Lower Murray, South Australia. Trans, Roy. Soc. S, Aust. 51; 

17M76. f 
STIRLING, E. C. 1911. Preliminary report on lhe discovery of native 

remains At Swanport, Rive Vi UlTil y: with an inquiry into rne 

alleged occurrence or a pandemic among the Ausir-dlian 

Aboriginals. Tranx Roy. Soc. S. Aust. 3S: 4-46. f 
TINDALE, N. B. 1926. Native burial at Pcdlcr's Creek. South 

Auaralia. 5. Aust. Nat. 8: 10. e 
1 INUAIlE, N. B. (atirib.) 1940. Skeletons at Brenda Station. S. Must. 

Nar. 20: 32. I 
WMfTE, S. A. 1919. Notes on the occurrence of Aboriymal remains 

below marine deposits ai the Rcedbeds, Fulham near Adelaide. 

Trans Hey. Soc S Aust. -43: 77-80 t 

(10) Dot mi: 

A. I. A S 1966, Reccni Avistralian radiocarbon dates. A J. AS MfWftA Mr 

2. 2O-20. u 
A. LA S. 1967 Recent Ausrintiau mcltocarbon dates. A.I.A.S, NkwfikfUifl 

2 20-3 3 w 
BARBLTTI, M., and KM ACH, 11. 1973 ANL r ( ,d.oo.rb<,n d;Uc hM 

V. KudiOLiJihnn 15; 241-251 £ 
BLACKBURN. G. 1966, Radiocarbon dates relating in .oil 

development, coaia-line changes, and v^lomiC Bsb deposition m 

south-east South Australia. Au\t. X Set. 19: 50-52 « 
BROLCKFK. W. S-, KULP, J. L M and TLCEK. C. S 1956. La«WU 

natural mcirocarhon meftiuittffncaffi hi Science 124 154-163 »' 
C*Ai-tOW, W. .1., pjQCj 'itherv 1964. National Physical Lalioraturv 

Radiocarbon tvltttsuremtnti fl. Razftorarbofi u. Z5»3fii t 
CAUOW, \v. J,, and others 196V National Physical Labonutity 

Radiocirbcn MeaMiivments III. Radiocurhon 1: l56,Jt5l t 
DLIRY, Ci. H. 1966/7. Australian geoclironolt.tiv. ^tirckltM'"!. 4ia/ 

./ SW, 2V: I5K 162. r, g 
I i.OOl^ J. |983. Appendix. Radiocaibon dales alerencc Ma. }n 

"Anrhaeology of'ihc Drearmime:' t.J. 1 loodj. pp. 23J-25& Collins, 


Cdl I FSIMI . k ( . and TEMPLE, R. B. 1977. Radiocarbon dating ol 
shell lttSd(ll!H& Arck Pftys. Ant/uop. Oceania 12: 26-^7. j; 

UlLLLSI'ljL, R.. jnaPOr ACM, JL A |976. Ihc sutiabiltiy of marine 
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»'~aliforniu, lulv 1976, r 

KKJOSHL K, and KOBAYASHl, H. I96V trakushin Natural Radio 

carbon Mejsuiemenis IV, kmUnrerboti 7: 10-23. g 
KKjOSMI, K. and others I%4 Caku'liin Natural Radiocarlu.n 

nllsaSUTemCfriLl 111 Radiocurhon 6: 197-207 « 
LAMPLR7. R L 1972. A carbon date for the Aboriginal occupauon 

,.l tfartgftrflp inland. South Australia. Maufyid W 22.V224 h 
MeC\RTH\, p. D 1966. A, ptissible Uvnd tf| r.idtocai^nn (Jal'e$ ffl 

Australia. AJAS •Vfewsfe/to' 2 27. \ 
MliLVANFY. IX J. 1959. Pflh'njai Australian piehtstor. Natiar 1X4: 

913 1 
MULV\NEV i\ J 1961. AuMuilian i.idioLarbon dates Anttniats^ 

.17-39. r 
MUI VANLY. O, .1. N63 Antiquity oi 'nan in Australia: pretn | i 

In ,l Au%trahan Aboriginal Studies: A synj|)odum!' (H Shuils, -A \ 

pp. J3.5I. O.U.K, .Melbourne. \ 
POLACH, H A., ftfld GOI SOW J l l >6.s- I oUcchon and suhirnsMo,, 

of radiocaibon samples, In "Ausirahan Archaetjlocy. A Guide 

to Field lechini|ucs M (D. I. Mulvancy, cd.), pp. 211-239. A-l ^ S-. 

Canberra- \ 
POLACH. M. S., and other-. I9J5&. \Nl ladiocarbort date lia II 

Radiocarbon 10: 179 19). u 


KI.1 'v v si \u H lv ,|> l u 

\ti/\: #tt 

PRLSCOl I, J ft,, POL AC H, H. A.TR! 1 IV CI I . and SMITH 
P. W P)XL Liamparison nl CM and thermoluminescent da.tes 
from R.-onka, Souih AuMralia B4C f JOU&WM* * 90-211, 1 

SHUTI FR, R ..It I97B, Radiocarbon itainf and Oceanic men- q 
.•1n77. and f'hvs. Atahmp Otraniu IJ: 2I5-2ZB, h 

i PMPLE, K. a and BARBLTTL M. I*»81. Rcvfttaiia m raJioveibon 

,l.iir-> mam ilieSvdnes L ra\t:rsHV Radiocarbon laboraiorv AUST. 
ARCUAEOL \h 2B » - 
ONDALL, N. U. I%4. RodisTCiirbon dates ol imereu Lu AusiialLnt 
aa'hacotoaw; dwf 7. o/ .vhmv 17. - ,J g 



Commmec -South Ausiialian sub-Commiiiee Report ch. 5. 

A.I.A.S. 1970, SiW5 m South Airstrali;* proclaimed IMlldrlhc Mxinynml 

and MKtoik Relics Preservation Act iy<<s. .i./..L\ s^wslvitet 

I ■ i ■ 
tOUTT^ P J. R l%6. Inierpietation ol aichacological tnalenal bom 

HiH'ace siies hi Australia A LAN, "Ivpesciipi. 

in Souih Australia^ Govt Printer, Adelaide. 
MOL STM\, r_ iy76. "Aboriginal material culture- of South AiUtttnOE 

A tvadinc tiutde" A.H.R.. Adelaide 
MITCHELL, S K. 1962. Analysis of tome AuMtuliau Abonimtal 

surface jilt'V. Mankind $: 466-470 
I'RF I J V, G, 1 L962. Archaeology in South Australia, llw Stitf: 7 


PRl ITVi O. L, iSl7(). Sutvc\ of prehiMoni. mo mis »• Sunlit 

Australia, AltSt. Ah. SfUd 11- }9 ••!). 
PRL M V. d I 1913. Archaeology in Souih AuMiaha -icpuil on 

recew wflrk, Atat Archacol. 3: J2-39. 
PRLTTV.C. L. 1977. Aivducolo^v in South Australia \A. \<ut IUn>k. 

I .o\ I Pnttlrv, 'Vlrkudr 

WARD, i, K., CAMPBftsI I , I ft, HALfc, H M., a..o TlNOALL, 
IN. li IM36. Fossil man in the state ol south Australia. M'/a Hash, 
Groi Cwtg Hu 1271-1273 

WEA1 ME KSHf-F, R. D, l%7, Surface collect iin- on StJUlll Australian 
L.nnp sites. 7. Andirop. Sue. S. AMff. 5: 7-10 

(a) IVe.stern, Nonh-Wesietti 

BASLDOW II Iyt4. Journal d ifte Government North West 

evpeditiom /Vrw /v'or CiflWfl -SVn-. 4wS/:. X l'/s<- /V/.///r// 15. 

IJROWN, M. I*77y- Mo.pholo^Kal irvunsiru. ».ou Cll (\nQ40AVlt.l^rfl 

Deseo. B *\. HOlm Ificib, - 1 - N I. 

-MiimirI . . undci cbniinand of IX lantlsav. Bmxuiw. Adeiiude- 
* Vk> . I J, 1X45 JourrtHkol B\pedhiionv of CJiacdwry intjj Cwiiral 

Atisimlia eic l ontiorv 2 

t ll/PAIRlt K. t 3 tSt"7 Survey of Ahoi.^iila! rfte« al.-iij ilu .., 

Of [fie piVpOiod I&tCOoIfi '" AliW Sprmes Rail's, i\ 
r 1 1 VI'ATKK k. R and OARA. I, 1984. Aj|throrro!psfc»| a.crv of 

CMf;tcti\e mineral leasee on Birthdav. lour Mile. BTTvl NauoUu 

Lreeks near fort Au^iMa. A re-putt loi die Kok.ittiu Kvpk . 

( otniiutlee AH H. (Reslr.Ued) 
CAI I L'.S, A. i%4, IWil Nustialiau ■■iiaii^raphic -.CMiicnves. f itrr, 

Amkfvjl ft 127. 
CiOUl U R. A. 1*>7|. T|« ardiaculuyra j|f BlIlRtJUllJijTCT ,i U|$e foil) 

|jf€ WfiblCm Deseri Off AustiMhw- World Anhoeuht^v JP; I43-T77 
'.oUl.n. R. A ls>7u_ [:sotic Stunei 'oilI l^aticred bonev cihnn 

archaeology in the AuslraUffri l>ev.'fi. Anlnwoh^v 32: 2fi-H 
t.OlH.D, K. a! I9H0. "IJvinji ArchaeolojA'' Cam. L Pr._*s\ New Voik 
t .Rl IN WAV, i IV73. "DOWii Atnonr The U -||d Men. The Nanaloe 

Kminatol Fifteeo V^spurMJina the Old Sittnt a^l Aborigines 

liI \uvLi.ilus WWem Dennr Htitehinson, Richmond, 
IOHNSON. J. E ii-d- Mamisertpt Of urcliaeoloyy ol S.W ceiilralhiu 

dcsefi. Sotehnoks. S.A. Museum AD 42, 
[OMN60N, J- i 1^53 Observflflorw oil Mime %bpr!gnml eamphites 

in South Australia and adjoining States. Mankind v M~79 
IDHNSON, J. l_. IV64. Observations on sonu XburifJUiiil LjmpsitLs 

in Soulh Ausiiaha and ad|oinin L ' SldU'J Wunktfttt (* 1^4-IS!. 
KOKATHA PEOPLED I OMMITTEE, 1983. John m, Trcyen/a. 

Kokailia Hei nave Sui\u\, Report un pilot stu-Jv. pCTDtao l'*MJ 

18 MmtlH 1083 A H R (Kcslrieled) fl 
TIMDALL N ft IW, ii rll bPltcfci III the Western DrMN Itl 

Australia Pan I— fthrrtOi 'a hu became the Pleiades, ftflp. V ^IwW 

V/^v IV Mi5-H2 
WHIII-, S A_ IW. "I,, lire t,r N.Hll.AVcst \n I v|V<la (0 i|„ 

MmgTave and L^etatd Ran-e--. Ihuma 1 . Adelai'le 

ib\ Fxtstern, North- Eastern 

MioRliilNAL AND HISTORIC RFLKS. |y?M Report ol >miJ J tiSTOrii: ReHcfi Lam. (Oepi. lui Lnvimnmont, S.A j 

ai HoiicvmiKxt ilepovil K;Pkartn» I»t Mines 

Admtnistralton reion. 
BECHTftl MNHIU JdlNI VI-NTL'RL. I9S|. Muomha io siun\ 

F'uiui. Dmfl F nvirouniental Impact Siatcmeni Report lo Hipc 

lines Autlioritv ol' S..\. 

Point pipeline faeihlies, neeUnalmn ol environmenlal FSki'Iun 

[<C|hhi lot Pipelines AulfimJiy \A S.A 
i ALLLiy R. A (eorni>.) lyso. Curiem icseareh itnhe I akohiome— 

' .iHahonnu atea. South Australia (includes ptclimiuaiv repoii h> 

) :!!'•-, I- 1 f.. and Lumpen, R-l.) -\u\L Quur \ind_ 14- 1207. 

C'OOPI IC H M I'^l Roek earvtnps and olhct Aboriyinnl te lies bom 

nr:u MaiTer V, \ini, V t ?/ 21; |-4. 
I INWARDS, R. [tffiS, Roek eugijvines and Ahoiiyinal nccu|Miion at spn ni's in dtc Northeast ol South Australia. /&v. i Attsi 

MUX 15907 
OAI* V T-. mi K* KJJHfcS, I J I I*ft3, An s.irvcv ot 

ilie i'iopo-,-..J < 'Ivmpic Dam livpass Road, s.A. \ tep<>n id the 

Highway) Depamn^i, Walkerville, S.A 
HI Wl 1 I. R. I''7^, | he -Veooiia I'latcau -ai\ outline ol ,\bonpiua! 

habitats and relicj. ./ Anthrnp. Sin. s Ahsi. Id: 6-iv 
HUOHI s P J WB, Aborietnal aichacoloetcal sites in lite Sainos 

I >cwiopmem Areas at RedclilT and m the <. OOPCI H;isin \ rcptefl 

i" Saiiivs I l<3 dirouglt National Sysiems Re-scarch h\. I td., I\m' 

Melbourne, Vie, fRestrieiedt 
[lUt.HLS, I 1 !. mO. Moomba RedclilT Pipelines L. I.S.: Ahoiunnal 

and Historic Sites A leport in kinnaiid Hill de Rohan mi til Vi ; 

Ph I Ul„ Melboutne. <Reslneled), 
Ml <..lli:s. P. J. 1081. An afchacDfdfiical invesii.uaoon nl ihe WUU 

ot the proposed road fa>m Phillip f'onds lo Purple Do\wi.. \ 

iCtHJrl tO Rchv Vk.iia yen lent Set wees Ph. I id , tltKnu-h Kinhdl 

[ J lv. 1 id., Adelaide. 
HiJCiHtS, I 1 - I- 1981 Olympic Dam Project A.eltaeoli^.eal Stutly 

ol' Wenan Shall Report to Ro\h> M;maLvuu-nr Snviees P|>. I, id., 

ihitnigh Kinhill Piv. Lid. 
HUCrHLS, R I 19X3. An -a.ivey ot iIk tcplacemeni 

Mooinha W iltoii ( ,.iv Pipdiue, SlrVeleeki Oe^eO, S. \. A report 

to ihe Pipeline Auihoriiv, Canberra. 

HI'OHI : S, P _L ( and HISCOCK, P. I9S0. \n andUPOtngiCfll 
Invevilyalion tff the route of ike proposetl road friffm Olympic 

I'iim Village to Purple Dowm., \ icport 10 Roxby Management 

Sei\K.e, P'\. I Id , AOeLiKL', S.A. 
III.<.HI\ P, J„ and HISCOCK, P. mi OKmpic Dam P.oi-.i. 

Itasehne \lctlOCO toyical Studw lor Kinhill Plv. lid. 
Ill ' IMES. R J-. HISOOCK. P, and KHOADS, J. I9H2 Oly.upn_ IXim 

Pii-jcci An.liaeoUjey Studv Ihe tyS2 Piojeet atea. HAvnsik and 
nc euriidot surveys inethodolcjjiy and results. A import >o 

KOKb) M.iM'iijru.enl Sen lees Ply I 'd ihiomili kmhlll Piy I id,, 

IILOHLS. P I., and LAMPORT, R. J. 1980. Pleistocene oeeupaimn 

ol the aiid /one in kOUll) easi Australia: reWKR.ll pio«-petis hu 

lite '. hiipu r "rcck-Sn/eiecki Desert rcpion. 
1I-MM -V A., and WILSON. A. IWO Kn tHVttO eutal Ptol.leSiudv 

01 the s< h ir i shell-t entral Tablelands Wc-ocm 

s..(Mipl,\iiii.ui. 1 M L'liiui Region "i South Ausimlia. 

Assessmetil Seelion, South Auslralrao Dcparlment lor Ihe 

I iiMiiuiineiit 

JONLS. W L. I"~'» i Iri \Ur eteek: huntei-piWercn in the Conpei 
Ha-aii li, \. Hon,. Mie-is, Univ. Nov\ I:it fe dand 

KIMHI I SU ARNS, vm, Olympic Own Project Supplement tu 

ihe Diah F*n\irt.Hinieutal Impact Slyiemem Prepated lui Ro\by 

M;ipuc.cmeui Services Plv. I id. 
MNHIU -STrAKNS JOINT VINllKl, P>82. Olympic Dam 

Ptoiect Dratt l-.tnironmeiiial Impaci Siatemeiii. i'repared ior 

Ku\hy Munae'i-iuem Semees Ply. Ltd. 

ILLMHI K>. R. I WOT Survav Rrpnrl I ItineymoLin lliamtim Drpnsii 
rtl Kalk;irnu S|,nioii \lJ01 n-iual 1 lerilaer I Init, Depi lot 
I nvuotiuicm, S.A. 

MAWSwN, D., and MOS.S1 I I D, P. S 1936. Relic-- o| .M-nnemal 
oioii'inun, Olat> Distnei Hun;, A'*a Sac. S. \a\t. 50: 17-2-1 

MeBRV!>P. I. lykC Kepon on the AiLhaeokteiLal leaiuicsiii stmie 
inNtholoei'-al and ni>U>rK mic< in the lake 1 yre Souih reeson 
Piepaietl Un Kinhill Ptv Lid in relalion to 1 be I llyHJpk l),.m 
I'-!-' I LS- 

\W< 1 INMJ ! I ■ A - ly?*. AtHMiemal Made m ihe I ake bva -cinon, 
\|.A Oual, thesis A.V.C. ( anbeira. 

NOHRS, M |97*J li|VCi)I0n Aboin'iual anil lift I ii 1 i. ' i 
Pi'uiiUai'o Nbitnw K'esei\.- ii R Lile. fRc^nictedK 

SMITH. M, A, pJ 7 y s.ieha'.'ol.<e'c:il o.^earvh on Plmnbajjo Hisioric 
RtiCJVe. )nnt: A<lniannithuh Tmfim Ai-nv/r/A.7 2 ft .H 


SMITH, M. A. 1980, SMttMli ttftyftlfne jWfrgV and Ktitaumi 

panem j s'.Mtin.HiL archaeological survcv ol PlUjfldftgC mjooh 
KkldttC ttrscrve, Somh VtMialu M.A. Thesis. A. N.L 1 - Canbcjru 

ill bUMAN, R- k. iSfcC). The atchucologieul IKftcffltii tfl ihc Coopu 
( nr': n-nnm (fcCp<Vt '•! .nluV^W- I Anthrtip. 'v.. v tiut 7: 5. 

I- >L< III K, II, M I, i":si). ippc CiiOPfci 'W l"r * Report). -/ 

(c) Mid-North, IVttders Kunges 


Pbim pipeline facilities bieddration or environmental factors. 

RCpOi Hpi.-lincz Authority of South Australia. 

DliCHTLL-KINHH I JOINT VfJNTLKt \ Nftl. Moombu to Stony 

Point. Dwfl FriMroiuuriiiat Impact Suiemeni ft& <rJ Id Pipelines 

AuihoriO. of S.A. 

i imbeivati Ahwigma] vew repon by Adnjarnirhnnhfl.n»neet 


Flintier. Ranges— An Abonuiiu.i Vic* A&tjt 

Minerawuiu, Ram Paddock Gate. An historic Adttjamaitianha 

settlement in * he* I tinders Range*. South Visit alia AH R. 
Til IS, R W. iy?5 The funeral practices and beliefs bl tlw 

Adniamalluinha / WtiMft Sim.. S. AuM. O: VIO. 
I I l/PAIKICh., P. I9H4 Preliminary anthropological mj> vcv ol 

Nalcanootia pornioil of ihe Gammon Range National TYnk, 

A. U.K. iRtstiiclcd). 
ioH vv, J |OTll No^GS on native tribe formerly tesidetu al Orromo, 

South Av»(ift|i4 $ AU$t Nor 12: 4-6. 
H.VCiFN, R. 1983, Repotl h> tfct WejflJI 8 Commurmv concerning 

the Beverley Uianium Project. A.H.R, (Rcslnc(cd). 
HOSk yns, ^. . iw6. Hie Hinders Ranges s.ic recanting pn>*ranimes 

1 Amhrop. Soc £ Aust 14: 6-10. 
Ml JC.HKS, P. J 19HU. Moomha-Redcliff Pipelines t I & Aboriginal 

and I lisiorie Sites Repot i to Kitm&ud Hill de Rohan and Young 

Plv. Lt4„ Melbourne. (Reslrictcd). 
ii ihnson. i, F |9ft3, observation-- im lorne Aboriginal campsites 

in South Austr&ka and adjoining states Mankind &(2). 64-791 
I AM PERT, R_ J. 1976- DempseyA lake palaeontoloeieal and 

archaeological niic-. ^1usr. A,vhoeul S: 12-14 
I AMPERT, R. J. and HUGHlrS, P .1 W<1, Plcisuxcne aidiaeology 

hi the Huidfis Ranges: research prospects. Ausl, ( 1n/?<vro/ 10; 

MARTIN, S. 198J Tooih Nuh Staonn AnhiieoloeiLjJ SUffi Atvriginal 

i-leritHge Section, Depi. lor Fiivtionmeni. S.A 
MINI. HAM. V- M. 19KA u t ho siory fll ihe flinders Ranges!' Rigbv. 

POriFK.I- 14X1 Pipeline facilities Moomba -Sionv Poini, Kep-m 

on the mipaci Of the f>rO|K^cd pipeline on Aboriginal ethno- 
graphic tilts, Neuroodla -Stony Ralwi, ikvliiel-Kinhill loint Venuire 

h>i rvxiMr. Aiubortt) of S.A. 

ROSS, B 1979, Sue Po«;iiiimiiniit;»n— Belunj. A.H R (Reslrieted). 
SNOFk, W, 1982. The archawlogv or ihc Ml Remarkable Naiionai 

Park. Wangvarra area. National Park and Wildlife Scvice. 

Deparnnrnt of Lnvironmem and PUnnin^, S.A 

Beverlev Project. Lnvironnienial Impaei Si.ucmem 

Heveilev Ptouxi. Dtafi t-PHroiM.eniui Ni.paa siatemenf. 

Supporting Document 
I LINBRIIXjH, 1984. Oocumcniaiion of Abouginal place uuroev 

Gammon Ranges National Park (Hid Siudy). A.H.R. 

MJKNFR, I, .mdOARA, I- i- 1982 275KV Transmission line. I\rn 

Ativitisla EQ AdHnule: Aaliao'lc^ical CUtVGy from Tiingkilln IQ 

Fuditntla- Report for Fleclricity Trusl of S.A, 

(d) Eyre Penttmda, Yorke Peninsula 

UK HtM-klNHll I JOINT VGNTliRt, 1 MM 1 Moomba. to -Stony 
Poini pipeline ('acililio. Heclarilton of en-. 'triHiiiieiitiil fueLor>. 
Report for Pipeline's Authority of South AuMralia. 

mCHTn-KlNHII I JOINT VbNTl'Rl:. P*«l. Moomba to Si&ny 
Point. Dran P'nviranmemal Inipaei Staierueni- Report 10 Pipe- 
lines Authority of S.A. 

I AM^HLiL. T CL, and WAI Sit, U. O. l*Ml Nflttl on Aboriginal ires on Yorkc Pernnsala, South Australia Mankind 3 

GARA. T I. IVS-. Archaeological survey of the |toOpb$60 PWlW (>l 
the Port Augusta io VVhyatla TrariMiii.ssion I itte. Ricpon o| Social 
and Bvofogtsal AssesHTiBni Pte i »d • tor Fiecmcnv trust d| s.a. 

liARA. 1 1981. Archaeological survey of iIk proposed route of A 
l33JtV irantfirisian line rr©w South Hiimmodc^ SutfiWifoo \u 
Kadina Suh>iiJtion. A.H.R. iRcsmetcd). 

HII I . 0. I... and Hill . C. J. 1975. Nutcs on the Naiutigga Tribe 

of Yorkc Peninsula. Lutheran Publishing House, Adelaide. 
UtSCOCK, P., and MCiuHES, P. J. ISftO. Aboriginal Aa'haeological 

Site k fn Soelj) 9fld F^»ln-h.ji .\-,essHu:nl Pty. I. [J D..O' 

F n\ iruniTivTilul Impaci Saiemenl lor Port and Terminal facilities 

at SlOlty Point, SA Ruckeround fttjPCre. Prcfta/cd t'oi Santos Ltd. 
M K, P., andHUC.HFS.P- J 1980. Abor.pinal archaeological 

sire: in the proposed Santos development areas at lA>n Snns'ac 

ftiKl St.Hiv PtaiQt. A l£pGn to Status ihrougti Social aiul Ucotogieai 

Assc^Mneni. Kern Town. S-A 
MUGHES, P- h I9K0. Aboriginal archaeological siles in the Santos 

Development Areas ai Redchfl and in the Cooper Basin Repon 

10 SlltlOS Lai. through Natural SysteirtB Research Ply I id- Port 

Melbourne, Vic. (Restricted/. 
HUCIHES, P. 1 I9S0. KedchlT Pipelines F.I.S AbonpinaJ 

and Historic SiCfei Report (o Kinndird Mill de YOUIg Pt/ , Melbourne. (Restricted). 
JACOBS. X. and POTTF.R. L I9KI. Stoir, Point Port and Terminal 

facilities— Anthropological Ass*:ssmcnL Report for Cooper BaMn 

l^ruducers LkfUSai Devutopment SchcuiL 
i lit FY. I A., and HcGHLS, P J. I9SI. An jRhaeologtcaUum-y 

of I he proposed Pipeline route from SlOlty Pomi ioCompressi-.r 

Station four A repon to Kinliill Ptv. Ltd., Norwood, S.A. 
MARITN. S 1983. Ticker.»AVallaroo Power Station Pioject baseline 

AnAaffOtagfcaJ Survty ol the proposed power siaiion area Repon 

to Km.ImII t-M Ltd AHR (Re:ancted). 
MARTIN, S I'JS4, Repon On thv BurVey ol fisl. irjpv di San CfWt 

iPorter Bay, Port lincoln|. A.H.R. (Reslnckdl, 
MOl'NTFORa C P. 1939 Aboriginal metWs of fishing ond 

CL'olanp a.-, used on the ftOiRhem CG3XI ol EvrcS Peninsula, South 

AUfcfraha Mi/nkmd 2: 196.200. 
PARSONS, A r |Q7|_ Am ahiHiv.MiiaU-yitip:ute on Yorke Peninsula 

f.O.S.A.M \ewsit-ner2: 4-5. 
PYRSONS, A. C 1979. Aboriginal sites on southtnt Yorkc Peninsula. 

/. 4tuh{,>/! W £ AuU- 17: ^-6- 
PGfTTER. 1 . P'HJ. Pipeline facilities Moomba-Stonv Point Report 

on the nnpact L«r the proposed pipeline on AOongiral eihnei- 

g^aphic sites Neunx^Ha -Smns 1'oini. Rw-htcl-Kinhill Joint Veriiuie 

for Pipeline AulhorHy. 
KOSS, B. I I9S2. Archaeological Survev, l?2RV r I ransmission line. 

Yadnane lu Wudinna. Mtd-Fyi^ V-'eumsula. SA- Report for 

Llectriciiy irust ol S.A 
KOSS, Q 19S3 tncnonmentAl impact Repon Heritage Survey— J*>TTer 

Bay. Prepared for the Department of Tourism, SA. AHR. 

ROSS, H\ I9K.i. I ineolu Nutioual Paik, An inventor of sues of 

historical significance a H R fRe»incte^) 

ROSS, Ek and JEFFF.RY, H. i9?4. Rcdcliff Impact Study. 

An jot jotogieal surv^Vi c^Dnuftdo report, prcrart^ toi the 

Peirochemicul ennvrttutn of Souih .-Vuslialia by ihc Aboriginal 

and Historic Relic:- Advisory Board, d 

Drafi Fnvimnmeiual Impact Slatcnient for Port and Terminal 

lacilidcs at Stony Point, S.A. Prepared for Santos Ltd. 
An TON, T M IK8W. J he AdjahdurJt Tribe o( Abonannes on Yotke's 
Peninsula Pro*. Of GM& S(«. ufAfiSU ^ Ausf - Onanch 17-19. 
1 1NDAI E, N. ti. KU6 oTQttt on rtaiives oi the souihern ponion of 
Yorke Pemii-->ulj, S.A Trans Hoy Son S Ausf. 60: 55. 

Fdiuanon, Sidn.hnry. 1981 Ihe History of Woman Habitarion 
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(e) Lower- North t Adelaide Region, Fieurieu fcninsuta 


S A MUSi'l Al, 1976 North Fast Area Public Transport Resiesv. 

Archaeolo^ieal Survey. Piclimutary Repon. 
CAMPBELL. V M. |979. Interim report on the Fieurieu FVninsub 

Projeti ./. Atulimp. Sioc S. Au.\t. 17: 14. 
CAMPBLLL, V M. iy79. Archaeological reconstruction Ol eoasiai 

sjies south o| Adelaide X Awhmp Sen- S. Aitst. 17" I 10. 
CAMPBEL1-. V. Mi 1980 C.vcavaiion of a small hcanh silcal Moana 

Beach, south of Adelaide. J. Amhrop. Sck: S. Aust, IB: 5-31 
I AMPHFI I . V. M I98L Archaeology of the souihern .Adelaide 

region. Introduction. J. Amhrop SOf, $ Ausi. 19: 1-5- 
t AMPHFU, V. VI 191st Archaeology ol the vouihcru Adelaide 

icgion e^n 2. Sitc-s and sequence*. ./ Anthrxyx Sac. 5. AUSt. 19: l-V 
COOPfcH, H. M. 1968. An Wthacolo^tcal camp- site in the vicinity 

of Hallclt ', o\c, Si»n'h Ausiniha. Pet:. S. AuSf. XUtx 15: «»-614. 
COOPER. PL M. 197e.. The record oi man In "Halkli Cove. A Field 

Guide" (H M. Cooper, M, Kenny, anil J_ M. Scrymgour. eds-K 

pp, 22 ??. S.A. Museum, .AUelaidt 
EDQUIST, A. G. 1^20. Naiutc'*. Wa\ (Aboriginal caniptne ground- 

lulbam, Adelaidet -v Am IStol- l:#W 


m . - VI i \u s. i9ll> i u 

WtfA flVW 

El WARDS, K. »9fi4- A former Aboriginal campsite on the Siun Rive 

al Marion. South Australia Mankind & 184*188. 
EDWARDS, R I96G. Aboriginal camp-silt' on the Sum River ai 

CnmmaiiJtrl V.dlev, South Australia /to.'. .V yfl0£ A/u.s. 15: 

EDWARDS* R 1972- "The Kaurna People of the Adelaide Plains" 

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REC. S. AUST. MUS. 19 (1): 1-14 

Maw 1985 

Aboriginal Archaeological Sites in South Australia 

P. Jonas 1985 


>10,000 years B.P 
<10.000 years B.P 
< 5,000 years B.P. 
Stone structures 
Rock paintings 
Rock engravings 

FIG. I. Map of South Australia showing the regions referred to in the text, Symbols denote localities of different subjects listed (see key 

for explanation). 


by Robert J. Raven 


A new species, Cethegus ischnotheloides, is described from South Australia. Males possess spine- 
like bristles on the palpal tarsi that are attenuate as in Ischnothelini. 





Queensland Museum, Gregory Terrace, Fortitude Valley, Queensland, 4006 
(Manuscript accepted 22 March I9S3) 


K Wf \. R, .1, |98.\ A new .ind interesting fcpa?tcs of CViiwmn 
Tliorull (UchnorheliiuiL- Oipluridael from Soulli Australia, AVr. 
H, Am WMfc \*H2): 15-1**. 

A new species, Cethegus ischnothetoides, is described 
from South Australia. Males possess spine-like bristles 
on the palpal tarsi that are attenuate as in Ischnothelini, 


Previously, the presence of spines on male palpal 
tarsi was known only for the western European genus 
Phyxioschaema in the Euagrini (Raven, 1981). However, 
only Ischnothelini (in the Dipluridae) have the 
characteristic extension of the male palpal tarsi (see 
Raven, 1983). In mos! Euagrini, palpal tarsi of males 
are short and truncate; in Cethegus ischnotheloides, 
the male palpal tarsi are \spinose and are slightly 
elongate apically (unlike other known Cethegus), 


Until recently, only three species of Cethegus were 
known from Australia (Main I960). In a revision of 
Australian Ischnothelinae, Raven (1984) clarified the 
diagnostic characters of Cethegus and C. fugax 
(Simon) and described eight new species. Main (1960) 
included specimens from eastern South Australia in C 
fugax, However, Raven (1984) included only specimens 
from southwestern Australia in that species. When 
Raven's study was completed, Mr David Lee, Senior 
Curator of Arachnids and Helminths at the South 
Australian Museum, presented me with two interesting 
males of Cethegus from central South Australia. One 
character of these males is unlike that of any other 
Australian ischnothclinid and requires amendation of 
the diagnosis of the Euagrini. 

Abbreviations are standard for the Araneae and with 
methods used may be found in Raven (1984). All 
measurements are in millimetres except eye measure- 
ments that are in graticule units, each being 0.025 mm. 


Tribe Euagrini Raven, 1979 


Diplurid spiders with long posterior lateral spin 
nerets with long apical segment; a hirsute pigmented 
cuticutar crescent surrounds the base of the posterior 
median spinnerets. No cuspules on labium or maxillae. 
Spines present or absent on normal or attenuate male 
palpal tarsi. Cheliccral furrow with one row o\ teeth 
on promargin Trichobothria with corrugifonn collar 
around bases. Tarsal organ low. 

Tribe Euagrini Raven 
Cethegus ischnotheloides n. sp. 
(Figs 1-4) (lable I) 

Males with elongate embolus reaching proximal 
palpal patella; spine-like bristles present on attenuated 
palpal tarsi. Sternum with blunt hairs. Females 

Holotypc male SAM N1981 394 

Carapace 6.56 long, 5.69 wide. Abdomen 5.38 long, 
4.13 wide. 

Colour in alcohol: Carapace and legs reddish brown, 
chelicerae maroon, Abdomen entirely brown, 

Carapace: Fovea short, semicircular, deep. Lateral 
margins with few weak lateral bristles. About 3 pairs 
of fovea) bristles. Striae deep, glabrous. Clypeus 0.18 
wide. Black hairs on inierstrial ridges. 

Eyes: On strongly raised tubercle; group is 0.27 times 
head-width, and is 1.59 times wider than long, Front 
row centres form strongly procurved line; back row 
centres form slightly procurved or straight line. Ratio 
of AME:AEE:PME:PLE, 18:14:11:13. Ratio of MOQ 
front wtdth:back width:length, 30;33:2K. Eye inter- 
spaces as AME diameters: AME-AMb = AI.E-PEE, 
0.3; AME-ALE - PME-PLE, 0.05. 

Chelicerae: With fine transverse dorsal ridges. 
Promargin with 4 large and 8 small teeth; basally wirh 
2 fine teeth. 

Maxillae: 1.80 long, 1.25 wide; with some short 
pointed setae; scrrula ridge-like. 

Labium: 0.63 long, 1.13 wide; groove broad and 

Sternum: 3.52 long, 2.92 wide; all sigilla oval; 
posterior and middle sigilla 0.18 long and away from 


kir s At si. \u s n<2): (5»|7 

Wit*: (MS 

FIGS. 1-4- Ct*the%us ischnotheloides n.sp, rmtlc holiMvpe. I. carapace and chelicerae. 2. sternum, maxillae and labium. 3. doisal palpal 
EArftft SncMtyg uucnuarion and spiuc-like bristles. 4. rcirolntcral palpal patella, libia and larsitff. Scale line - 2 mm for iig I; 1 mm 
lor figs 2, 4; 0.5 mm tor lig. 3. 

margin; anterior sigilla 0,20 long and 0,25 away from 
margin. Setae long erect or short blunt. 

Palp; Bulb pyriform; embolus long, tapering, 
extending to proximal patellae: tibia with slight ventral 
groove; tarsus slightly pointed with cluster of slender 
spine-like bristles. 

Legs: Leg formula, 4321. I^g I smooth, cylindrical. 
Tarsi 111 and IV erect hairs forming weak scopula 
divided by setae. Spines. No true spines on femora or 
patellae. Leg 1 tibia, pi v2; metatarsus, p2 v2; tarsus, 
v4. Leg 2: tibia, p2 v3; metatarsus, p3 v5; tarsus, v5. 
Leg 3: tibia, p3 di r3 v6; metatarsus, pi I r5 \8; tarsus 
d3 v7. Leg 4: tibia, p4 r4 v7; metatarsus, p9 d3 r5 vfij 
tarsus d4 v9. Palp: 0. Claws: 12 fine teeth on paired 
claws; 3 fine teeth on unpaired claw. Trichohothria: Two 
rows, each of 10, on tibiae; about 10, in a line on 
metatarsi; 8, of different length on tarsi in irregular row. 

Spinnerets: Posterior medians 1,20 long, 0.36 in 
diameter and 0.68 apart; crescent of cuticle present as 
darkened area forming an anterior fold. Basal* middle, 
apical and total articles of posterior laterals, 1.92, 1.64, 
2.04, 5.60 long respectively. 

Material examined: Holotype male (N 1981 394), 
paratype male (N1981395), sand dunes. Commonwealth 
Hill Station, 29 1 '57 'S., LM'MO'E., South Australia, 
April, 1981, P. Bird, R, Sinclair, deposited S.A.M. 
Disiribution; Central South Australia. 




Leg I 

Leg 11 

Leg III 

Lpg IV 




5 06 







28 1 



1 ibia 
















3 75 





20 25 



Remarks: Males of Cethegus ischnoiheloides differ 
from those of C, Jngaxm Ihe 'spinose' pointed palpal 



Inasmuch as maLes of C. ischnoiheloides possess 
'spinose' pointed palpal tarsi, they differ from males 
of all other Australian Euagrini and bear some 
resemblance to male Ischnothelini. However, because 
spinose male palpal tarsi are also found in genera of 
Masteriinae and Diplurinao, the presence of spines may 
be regarded either as a symplcsiomorphy oi a 
parallelism— a hypothesis involving their 
synapomorphy is falsified by the absence of euspulc.s 
and the presence of corrugiform trichobothrial base 
collars in Euagrini. J here regard the presence of spinose 
palpal tarsi in Ischnothclinae as a plesiomorphic 
retention. However, parsimoniously, the attenuated 
palpal tarsus of males of C ischnoiheloides is 
presumably a parallelism otherwise Cethegus and 
Jschnothele are paraphylctic. 

That a Western Australian cuagrine should retain 
characters otherwise present only in Indian and 
Neotropical (Ischnothele) and Ethiopian {Lathrothele 
and Thelechoris) genera is probably predictable. Some 
geologists (e.g. Powell el af, 1^81) propose that pre- 
drift India was in close contact with Western Australia. 
If that were so, some sharing of taxa dating to that 
period is to be expected. Spiders of the genus Cethegus, 
as defined by Raven (1984), are predominantly 
xerophilie although a number of species occur in 
northeastern Queensland, Throughout its range, to my 
knowledge, Cethegus is allopatric with other euagrine 
genera. 1 suggest that Cethegus represents a group that 
was first isolated in Australia as the most plesiomorphic 



genus (that proposition will be discussed more 
completely elsewhere) and as such has retained 
characters of the presumed sister group of the Euagrini, 
the Ischnothelini. Therefore, Cethegus ischnotheloides 
may be the most plesiomorphic Cethegus species. 


I am grateful to Mrs V. E. Davies and R. Molnar 
for commenting upon this manuscript, to Mr David 
Lee for bringing the spiders to my attention and to Ms 
R. Owens for carefully typing the manuscript. 


MAIN, B. Y. I960. The genus Cethegus Thorell (Mygalomorphae: 
Macrothelinae). J.R. Soc. West. Austr. 52: 9-11. 

POWELL, C. McA., JOHNSON, B. U, and VEEVERS, J. J. 1981. 
Gondwanaland, the separation of Australia and India and their 
interaction with southeast Asia, 17-29. In Keast, A. (ed.). 
Ecological biogeography of Australia. Monographic Biologicae 
41. London: W. Junk, The Hague. 

RAVEN, R. J. 1981. The mygalomorph spider genera Phyxloschaema 
Simon and Stenvgrocercus Simon {Ischnothelinae: Dipluridae: 
Araneae). Bull.' Brit, arach. Soc. 5(5): 225-31. 

RAVEN, R. J. 1983. Notes on some mygalomorph spiders in the 
Musee Royal de l'Afrique Centrale (Dipluridae, Hexathelidae, 
Ctenizidae: Araneae). Rev. zool. Aft 97: 549-562. 

RAVEN, R. J. 1984. Systematics of the Australian Curtain-web 
spiders (Ischnothelinae: Dipluridae: Chelicerata). Aust. J. Zool. 
Suppl. Set: 93: 1-102. 






byC.A. W. Jeekel 


Two new species of the genus Heterocladosoma Jeekel, H. zebratus and H. galaxias, and one of the 
genus Somethus Chamberlin, S. grossi, are described. Australiosoma castaneum Attems is 
redescribed and made the type-species of a new genus Oncocladosoma, which is probably related to 
Somethus, Dicladosomella Jeekel and Phyllocladosoma Jeekel. A new subspecies, O. castaneum 
ingens, and two new species, O. conigerum and O. clavigerum, are added to the new genus. The 
holotype of Polydesmus innotatus Karsch, a subadult female, has been re-examined; it may be 
referable to a genus close to Otoplacosoma Verhoeff (tribe Antichiropodini). A list of the known 
South Australian Paradoxosomatidae is given, and some distributional patterns are briefly 
discussed. Brachyiulus lusitanus Verhoeff" a west Mediterranean julid, is recorded from a number 
of South Australian localities. 





Institute of Taxonomic Zoology (Zoological Museum), University of Amsterdam, P.O. Box 20125, 1000 HC 

Amsterdam, The Netherlands 

(Manuscript accepted 22 September 1983) 


UtKLL. C, A. W, 19S5. Millipede 1'rum Auslrulm, y. 
Ausiuitiosonuuini from Souih Australia, ft it 1 1 n note on the stiilus 
ill My/leSttiU\ mtlolalits kjtrsch wit) I ir ^i rccuitl nl n slvdihJ 
Mediterranean iiilitt in Aus(r;ili;t (DiplopoUn: PolyilcMTmii, 
Pji.uloMisvuiniliil.KA: luliUu, lulidm), HtC, S, \hm. ,\/u.y I9M>: 
19 .17. 

Two new species of the genus Heterocladosoma Jeekel, 
H. zebratus and H. galaxias, and one oi the genus 
Sometlnts Chamberlin, & grossi t are described. 
Australiosoma castaneum Attems is redescribed and 
made the type-species of a new genus Oncocladosoma, 
which is probably related to Somethus, Dkiadosomeila 
Jeekel and Phyllocladosoma Jeekel. A new subspecies, 
O. castaneum ingens, and two new species, G conigerum 
and O. clavigerum, arc added to the new genus. The 
holotype of Polydesmus innotatus Karsch, a subadult 
female, has been re-examined; it may be referable to a 
genus close to Otoplucosoma Verhoeff (tribe Antichiro- 
podini). A list of the known South Australian Para- 
doxosomatidae is given, and some distributional 
pattens are briefly discussed, Ihvchyiutus lusnanus 
Verhoeff, a west Mediterranean julid, is recorded from 
a number of South Ausiralian localities. 


This is the second of two papers treating the 
ParadoxosomaJidac of South Australia. The first paper 
dealt with the Aniichiropodini (Jeekel 1982a) and the 
present conlribution treats the Australiosomatini of the 
State. As pointed out already, our knowledge of the 
millipedes of Sourh Australia is extremely scanty. In the 
family Paradoxosomatidac only two species had been 
described: Polydesmus (Strongylosoma) innotatus 
Karsch, 1881, an unrecognizable species of unceilain 
taxonomic status, and Austruliosomu castaneum Attems, 
1944, a reasonably well characterized but as yet not 
correctly classified species (ef. Jeekel 1968: 26). 

The material reported upon was received on loan from 
the South Australian Museum, Adelaide. It had been 
collected mostly in the surroundings of Adelaide, and 
only lew samples from elsewhere were available. 
Nevertheless, it gives a fair first impression of the 

composition o\' the fauna, and indicates that South 
Australia has a relatively rich representation of 
Paradoxosomatidac (together with the suborder 
Cambalidea of the order Spirostrcptida the family 
appears to be the dominant millipede group) with still 
many new species to be expected. 

The type-specimen of Polydesmus innotatus Karsch 
has been re-examined, but it is a subadult female, which 
cannot be associated with any of the other South 
Australian paradoxosomatids at hand. Neither can it be 
referred with certainty to cither of the two relevant 
Australian paradoxosoruatid tribes. 

It has been pointed out earlier (Jeekel 1981; 20) that 
records of millipedes introduced into Australia from 
elsewhere arc few. The material iti the South Australian 
Museum contains, besides some specimens of 
Ommatohdus moreleti (Lucas), a species now known to 
have become widespread in a large part of South 
Australia, also a number of samples of Brachyiu/us 
imitanus Verhoeff, a Mediterranean julid known to have 
synanthropc tendencies, but as yet recorded only a few 
times from outside the palcatctie region. 

The bulk of the material treated, including the type* 
o( the new species, is preserved in the South Australian 
Museum, some paratypes and voucher specimens have 
been retained by the author for the Zoological Museum 
at Amsterdam. 


Heterocladosoma Jeekel 

Heteroctadosoma Jeekel, 1968: 144 

Type-speaes: Eusirongylosonut hifalcatuni Silvesiri, I89S 


This genus was proposed for three species occurring 
in the coastal region o\ Queensland. The quite 
unexpected discovery oi 1 two new species in South 
Australia extends the range of the genus considerably 
and establishes an important faunistie link between i he 
fauna of South Australia and that of the Australian east 



5. At ST. Ml S. I<; (J); |t».17 

A/ (f >. /'AS". 

Heteroctadosoma is well characterized by two 
libiotarsal branches arising from the base of the 
acropodite of the gonopod. one narrow lanceolate, the 
other more voluminous, broadly laminate and apicaliy 
more or less recurved. The femora) process is coalesced 
with the solcnomerite over most of its length, its free 
part arising from the distal part of the channel-bearing 
branch of the gonopod as in Somethus Chamberlin. But 
contrary to Somethus. which has only one libiotarsal 
branch, the spermal channel does not make a loop, but 
runs straight towards the apex of the solcnomerite 

Hetetoctadosoma zjebratum n. sp. 


Coralbignie to Buckleboo, Sourh Australia, 
I5-16.U1.1950, leg. G. F Gro$s,<? holotype, 3 ? paratypes, 

North of Kokotha, South Australia. II VI 1956, burnt 
out of spinifex, leg. G F. Gross, 1 $ paratype. 

Nonning, Gawler Ranges, South Australia, 17. VI 
1956, burnt out of spinifex {fruxlia sp.). leg. G. F. Gross, 
1 $ paratype. 


CoiouK Probably somewhat faded. Head with 
clypeus. frons and anterior part of vertex brown; 
remainder, including lateral scJerites brownish yellow. 
Antennae light brown, infuscate towards apex of 6th 
antennomene; 7th antennomere black, its tip whitish. 
Intersegmental membranes of antennae brownish yellow. 
Collum with a broad zone along anterior and lateral 
margins brown, remaining part brownish 
yellow. Subsequent somites with a rather narrow zone 
of the prosomites adjacent to waist, the waist itself, and 
the pan of metatergites adjacent to waist, about hallway 
towards the transverse furrow, brown, fading below the 
level of paranota to brownish yellow. Remaining parts 
of pro- and metasomites brownish yellow. Venter and 
sternites brownish yellow. Legs brown, with pale 
intersegmental membranes and a pale tarsal tip. Anal 
somite dorsally brownish yellow, sides brown; margins 
yellowish. Paraprocts brown, with brownish yellow 
margins. Hypoproct yellowish, 

Wtdffcf: 2.5-3.5 mm, 9; 3.2-3.5 mm. 

Head atui antennae- Labrum widely and rather weakly 
emarginate. Clypeus rather strongly impressed towards 
Sabrum and with an impression on each side below 
antenna! sockets. Lateral border widely convex, weakly 
emarginate near labrum. Surface uneven due to the 
presence of setiferous pits. Pubescence moderate, setae 
of moderate length. Lateral parts of head almost 
Hairless. Frons not demarcated from clypeus or vertex, 
sparsely setiferous. Antenna! sockets separated by 1.5 
Times diameter of a socket or by 0.7 times the length 
of 2nd amennomere. Postantennal groove rather deep 
and wide» the wall in front moderately prominent 
Postantennai bean-shaped area indistinct, weakly 

demarcated and not inflated. Vertex longitudinally 
widely convex, mote strongly so near collum; 
transversely faintly concave, and laterally rather strongly 
convex, but wiihout inflated lateral edges. Venigial 
sulcus moderately impressed, running downward to 
upper level of sockets. Vertex hairless. Antennae rather 
long and slender, weakly clavate, with 5th and 6lh 
antennomeres thickest Antermomeres subcylindrical, 
but 5th and 6th a little more obconical; 6th antennomere 
not inflated. Pubescence moderate in proximai 
antennomeres, becoming rather dense in the distal ones, 
Relative length of antennomeres 2 to 6: 0.95, 1.00, 0.95, 
0.95, 0.90. 

Collum: About as wide as head, subtrapezoidal in 
dorsal outline. Anterior border straight in middle, widely 
rounded more laterally and straight again towards lateral 
sides. Posterior border widely and weakly concave, 
laterally rather weakly convex. Lateral sides almost 
evenly and rather narrowly rounded. Marginal rim 
laterally a little incrassate, not brimtike. Premarginal 
fmrow distinct, vanishing towards middle of anterior 
border. Surface smooth, hairless, transversely widely and 
evenly convex irj middle to become somewhat more 
strongly convex laterally; longitudinally widely convex, 
a little more strongly so near anterior border. 

Somites: Rather weakly constricted. Prosomires dulled 
by a fine cellular structure. Waist narrow, rather sharply 
demarcated from pro- and metasomite, dorsally 
distinctly beaded down to level of paranota, weakly 
striolate along sides. Metatergites smooth, shiny, 
hairless. Transverse furrow present from 5th to 17th 
somite, weakly indicated on 4th and 18th somites 
Furrow rather well impressed, with a vague sculpture, 
disappearing laterally at a distance from dorsal furrow 
of paranota about equal to dorsoventral diameter of a 
poriferous paranotum. Sides smooth in general, but up 
to 4th somite somewhat granulosa Pleural keels up to 
3rd somite represented by rather distinct curved crenulate 
ridges, concavity upwards, without posterior lappet. 
Pleural keels in 4th somite vestigial, in 5th absent. Sixth 
somite with a weak posterior swelling and 7th wilh a 
low conical swelling near posterior margin of somite, 

Paranota: 2nd somite a little wider than collum. In 
dorsal aspect anterior border rounded and slightly 
shouldered at base; latero-anterior edge narrowly- 
rounded, without distinct lateral tooth. Lateral border 
widely and almost evenly rounded. Latcro-posterior edge 
subangular, narrowly rounded, a little produced caudad 
and projecting weakly behind margin of somite. 
Posterior margin very short and a little concave In 
lateral aspect upper side straight, sloping a little in 
anterior direction. Marginal rim rather thick, a little 
callous, upper furrow distinct, also along anterior and 
posterior margins. Third somite a little wider than 2nd 
and as wide as 4th. Paranota of 3rd somite in dorsal 
aspect widely and evenly rounded, a little more narrowly 
rounded anteriorly. Latero-posterior edge angular and 



a little produced caudad, but not projecting behind 
margin of somite. Posterior border short, straight. In 
lateral aspect upper margin straight, curving abruptly 
dorsad anteriorly. Dorso-ventral width not much larger 
than that 6f paranoia of 2nd somite; ventral 
demarcation by a distinct impression, reaching forward 
to about three-fifths length of mctasomite. Paranota of 
4th somite rather similar to 3rd. In dorsal aspect more 
widely curved than in 3rd somite, with posterior edge 
subangular, not produced caudad, and posterior border 
straight. In lateral aspect upper margin curved dorsad 
more widely. Dorso-ventral width as in 3rd somite, but 
ventral demarcation reaching cephalad to about hallway. 
Paranota of 5ih somite rather weakly developed and 
rather weakly prominent. Margin in dorsal aspect widely 
convex. Posterior edge subangular in porclcss somites, 
narrowly rounded in poriferous somites, In posterior 
somites posterior edges become minutely angular and 
produced a little caudad from 14th somite onwards, 
projecting only a little behind caudal margin of somite 
in 17th and 18th somites. In lateral aspect dorsal 
demarcation of paranota weakly concave anteriorly, 
weakly convex posteriorly in poriferous somites, about 
straight in poreless somites. Dorsal furrow anteriorly 
niiher abruptly eurving upwards, but not reaching waist. 
Paranota dorsovcnl rally not wide, rai her narrow, 
especially in poreless somites. Ventral demarcation by 
a depression reaching cephalad to about halfway or two- 
fifths length of meiasomtte, Posterior edges in lateral 
aspect acutely angular, especially in poreless somites. In 
poreless somites ventral demarcation a little concave. 

Slcmiies and le&s: Stci niles of middle somites longer 
than wide (ratio 1.3:1.0). Cross impressions well 
developed; longitudinal impression raiher wide; 
transverse impression also rather wide, but narrow 
between coxae. No sternal cones. Pubescence rather 
dense, especially near bases of coxae; hairs of moderate 
length. Siernite of 4th somite rather broad, rather 
densely setiferous, and transversely rather widely 
concave. Siernite of 5th somite with a broad process 
arising between and slightly in front of anterior coxae; 
at base the process is a little wider than distance between 
coxae. Process short, projecting downward, widely 
rounded. Anterior side a little concave with dense brush 
of short setae, Posterior side faintly convex with some 
long hairs. Transverse impression deep. Posterior part 
of sternite excavate, but not down to level of metasomal 
filtgj not taiscd at base of coxae. A group of long setae 
in the middle, Siernite ol 6th somite deeply excavate; 
posteriorly level with metasomal ring but anteriorly a 
little raised above level as in posterior part of siernite 
of 5th somite. Coxal bases not raised, but coxae of legs 
dMincMy elongate. Transverse impression wide and 
shallow. Four areas with long setae arranged in a square. 
Stemile of 7th somite with a low and rather narrow, 
finely granular ridge laterocephalad of gonopod 
aperture. Siernite of 8th somite excavate, particularly in 
anierior half, and raised only a little above ventral level 

of metasomal ring. Transverse impression weakly 
developed. Legs rather long and slender, prefemora 
rather convex dorsally. Pubescence on ventral side rather 
dense in all podomeres, Hairs moderately long, Tarsi 
pubescent on all sides. Femora almost straight. Tibial 
and tarsal seopulae present on anterior legs but soon 
thinning out and absent from, legs of 7th somite 
onwards. First leg strongly incrassate. Co*a of 2nd leg 
with a short medial rounded cone. Relative length of 
podomeres 2 to 6 in middle somites; 0.65, 1.00, 0.60, 
0.55, 0.75. 

Anal somite: Dorsal profile straight or faintly convex. 
Sides of epiproct eoneavely converging, before apex a 
slight indication of an abrupt stepwise narrowing, quite 
near apex. Apex of epiproct with lateral edges narrowly 
rounded and posterior margin weakly concave. Epiproct 
broad at base, distally still rather broad. Dorsovcntral 
width moderate, length moderate. Setae not on tubercles. 
Paraprocrs with narrow, moderately high rims. Setae not 
on tubercles. Hypoproct large triangular, paraboiically 
rounded with sides convex and apex more narrowly 
rounded. Setae not on tubercles. 

l-ICi. I. tletL'trKliittoSinno Zt'hra/t/tH n sp.. hotolypL-J, nt>lu gOMflp&A 
mcjiyl flSpVVl 

Ganopocis; (I ig. t) Coxa somewhat elongate, rather 
stout at base, but narrowing distad. Setiferous area 
ralhcr large. Prefemuj short, ovoid; its distal 
demarcation transverse on axis of acropoditc. Tibiotarsal 
branches both well developed; caudal one lanceolate, 
widening a little distad; anterior one stouter, more 
irregular in shape, and ending in an acuminate 
somewhat uncate apex. Solenomerite well developed, 


'■:l ■ S ALSI \1I,K W**> IW1 

AA/i. rms 

with permal channel running along media! side, widely 
curved and apically narrowing to solenomerue proper, 
which curves a little mesad. Femoral process arising 
quite near apev of solenomerite, curving caudad, about 
as long as solcnomerite proper, apex acuminate, 

Female: Sternites as long as wide. Pubescence oflegs 
and sterna less conspicuous than in male. Legs shorten 
prefemora not incrassate. Relative length of podomeres 
2 to 6 in middle somites: 0.80, 1.00, 0.50, 0.50, 0.85. 
Head with vertex transversely more evenly convex, not 
particularly flattened in middle and without lateral 
swellings. Collum with anterior border much more 
evenly convex and only a little more strongly rounded 
laterally. Pleural keels of 2nd somite produced into a 
long process; of 3rd somite in a long posterioi cone; 
in 4th somite only a minor cone near caudal margin. 
Coxa of 2nd leg with a dagger-like process arising from 
ecto-caudal side of apex, pointing laicro-vcntrad. 
Epigynal structure consisting of two paramedian wide 
emarginations separated medially by a low conical 
process pointing cephalad. 


The three previously described species of 
Heteroetadosoma, viz H, bifakatum (Silvestri, 1898), H 
transversetaentatian (L, Koch, 1867), and 11 
hatnuligerum (Verhoeff, 1924), are all from the coastal 
region of Queensland between Cairns and Brisbane. 
Compared to H. zebrarum and H. galaxias these three 
species are different in that the largest tibiotarsal branch 
projects a little distad of the seminiferous branch. H, 
hamuligerum is distinct in having rhe seminiferous 
branch apically divided into three processes, a 
solenomerite and two, or possibly one, deeply split 
femoral branches. In ff. transversetaentatum and H. 
bifakatum the bifurcation of the seminiferous branch 
into solcnomerite and femoral process is situated eJoser 
to the base of the branch, resulting in a relatively greater 
length of the two disral processes. H, 
transversetaentatum is particularly distinct in its 
voluminous gonopod telopodite and in the apex of the 
larger tibiotarsal branch which is strongly recurved 
forming an elongate spinelike production. As yet it is 
not clear to which of the three Queensland species H. 
zehratum and H. galaxias are most closely related, but 
on account of the total configuration of the gonopods 
these new species seem to approach H. bifakatum more 
than the others. 

Hetemciadosoma golaxias n. sp. 

Gammon Plateau, South Australia, 19.JX.I956. kg. 
G. F. Gross, 6 holotype. 


Colon/: Head dark brown, with labial area, an area 
around antennal sockets and sutures of lateral sclcrites 

yellowish brown. Antennae dark brown, with 6th and 
7th antennomeres darkest; intersegmental membranes 
yellowish, rip whitish. Collum blackish brown, with a 
pair of semilunate yellowish white spots at caudal 
margin, almost touching medially and continued 
laterally in a yellowish streak along caudal margin, 
disappearing near lateral rounding. Somites blackish 
brown, ventrally paler. Sternites and legs rather dark 
brown. Dorsum marked by two series of paramedian 
oval yellowish white spots, occupying the metatergites 
from halfway between waist and transverse furrow, and 
prosomites of the next somite to about halfway to its 
waist. I .ateral sides a shade paler brown in their posterior 
third. Anal somite dark, but ventral side including 
hypoproet paler brownish, epiproct entirely yellowish 

Width: 3,3 mm. 

Head and antennae: As in the preceding species, but 
differing in antennal sockets being a little closer to each 
other, and separated by 1,35 times diameter of a socket, 
or by 0,6 times length of 2nd antennomere. Antennae 
with distal antennomeres a little shorter; reJativc length 
of antennomeres 2 to 6: 0.95, 1.00, 0.95, 0.85. 0.75. 

Collum: Differing only in being a little wider than 

Somites: Waist rather narrow, distinctly demarcated 
from both pro- and metasomites* dorsally not beaded, 
but faintly longitudinally striate down to level of 
stigmata. Pleural keels tn4th somite weakly developed. 

Paranota: Posterior edge of paranota of 2nd somite, 
not eaudally produced, posterior border obsolete, In 
lateral aspect the upper margin curves a little upward 
eaudally. Posterior edge of paranota of 3rd somite 
produced a little and projecting slightly caudad of 
margin. Posterior edges of paranota of 5th and 
subsequent somites narrowly rounded in dorsal aspect, 
becoming subangular in 14th somite only, not produced 
caudad. Ventral demarcation of paranota reaching 
cephalad to about three fifths of length of metasomhe. 
Posterior edges in lateral aspect rather narrowly truncate 
in poreless somites, more widely and more obliquely 
truncate in poriferous somites. 

Sternites and legs: Similar to those of the preceding 
species. Sternal cones are weakly indicated in sternites 
of middle pari of body. Sternite of 5th somite with 
process directed downward and a little cephalad, 
projecting scarcely in front of the sternite. Process rather 
short, its apev quite widely triangular, medially rounded. 
In lateral aspect posterior side of process straight, with 
a transverse concavity at its base. Legs rather long and 
moderately stout. Relative length of podomeres 2 to 6 
in middle somites. 0.50, 1.00, 0.55, 0.55, 0.70. 

Anal somite: Epiproct almost parabolically rounded, 
apex rather narrowly rounded, medially straight and not 



FIG. 2. Hfiennhhiosomu gaUtxiits n. sp., holotype^, righl gonopod, 
medial aspect. 

Gonopods: (Fig. 2) Largely similar to those of the 
preceding species, but anterior tibiotarsal branch quite 
different; narrow at base, gradually widening distad, 
sharply crooked, and curving rarher abruptly distad 
again; terminal part rather voluminous and tapering 
towards apex and somewhat uncate. Posterior tibiotarsal 
process distally widening, then rather abruptly 
narrowing and ending in a narrow spinelike process 
(strongly reminiscent of a similar structure in 
Slreptodadosoma dissimile Jeekel (1980: 8, Figs 8-9.) 

Female: Unknown. 


Although this species is obviously closely related to 
H, zehratum, it is easily distinguished by its very 
distinctive colour pattern, by differences in the structure 
of the gonopods, and some other features like the 
different sculpture of the waist, the slightly stronger 
development of the pleural keels, the less prominent 
paranoia, and the presence of weak sternal cones. 

Somethus Chamberlin 

Somethus Chamberlin, 1920: 651; Jeekel, 1968: 27; 
Jeekel. 1979: 651. 

Type-species: Somethus fusdpes Chamberlin, 1920. 


This genus was erected by Chamberlin for the 
reception o\ a single species from an unknown 
Australian locality. Unfortunately, the original 
description was not accompanied by a drawing of the 
gonopod of the type-species. On that account the 
placement in the tribe Australiosomatini was somewhat 
arbitrary (Jeekel 1968), A re-examination of the type 
material (Jeekel 1979) confirmed the placement. 
However, with regard to the gonopod structure, doubi 
still exists on the true identity of the species. 

The discovery of some species of the genus in 
northern Tasmania and Victoria, to be published 
elsewhere, and the new species described hereunder has 
proved that Somethus is a well-defined genus 
characterized in particular by having a single tibiotarsal 
branch, arising from near the base of the acropodite of 
the gonopod v a small femoral process emanating from 
the channel-bearing branch of the gonopod more or less 
near its apex, and a short solcnomeritc proper, having 
a triangular additional process, in which the spermal 
channel makes a loop before running towards the ape* 
of the solenomerite. 

The record of Somethus grossi n, sp. from South 
Australia extends the distribution westward into the 
moister parts of that State. It seems likely that the type- 
species may be found sooner or later somewhere in 
southeastern Australia also. 

Somethus appears to be closely related on the one 
hand to Phyl/ocladosoma Jeekel, 1968. and 
Dicladosomella Jeekel, 1982, in which the femoral 
process is completely lost and the tibiotarsal branch 
more broadly expanded, and on the other hand to 
Oncodadosoma n. gen., in which the femoral process 
is vestigial or absent and the tibiotarsus is also more 
voluminous and more or less clubshaped, The four 
rnenlioned genera arc furthermore each characterised 
by a particular size and curvature of the solenomerite 

Somethus groxsi n. sp. 


Rankin Creek, near Melrose, South Australia, 
13.V1.J954, leg. G. F. Gross (E&T. 1355), <t holotype, 
1 ? paratype. 

Same locality date and collector (E.S.I. 1335), 3# 


Colour: Head brown, with labral area, an area around 
the antenna! sockets, and lateral sclerites of head 
yellowish. Antennae yellowish to brown, with 6th and 
basal part of 7th antennomere darkest. Collum brown, 
with traces of a rather narrow, medially widest, 
transverse band along posterior margin. Somites also 
brownish, with most of area behind the transverse 
furrow, including posterior half of paranota, yellowish. 


R| i > uisi \)t;s Wn»)i "'' P 

Wc/v. tatf 

Venter, sternites and legs yellowish. Ana! samite brown, 
but most of epiproct, and margins of paraproets, and 
hypoproct yellowish. The material appears to have lost 
much ol its original colour and I he brown colour may 
have been blackish brown in living specimens. 

IVhitlvJ: 3.2-3.4 mm,?: 3-7 mm 

Head ami antennae: Labral emargi nation of moderate 
width and depth- Clypeus very strongly impressed 
towards lite labrum, the impression ercseentie in shape, 
and surface above it a little inflated. Clypeus with 
setitcrous pits, moderately densely pubescent; on each 
side below antenna! sockets a rather weak impression. 
Setae rather short. Lateral border of clypeus widely and 
weakly convex, a notch near the labrum. Literal sderites 
of head hairless. Prons not demarcated from clypeus 
or vertex, smooth, hairless. Antennal sockets separated 
by 1 .6 times diameter ol a socket or by 0.7 times length 
of the 2nd antennomere. Postantennai groove rather 
deep and moderately wide, wall in trout rather 
prominent. Bean-shaped area at posterior margin of 
antennal sockets rather distinctly demarcated, slightly 
inflated. Vertex almost evenly convex longitudinally, 
slightly more convex near collum; almost flat or even 
a little concave transversely, lateral edges rounded and 
somewhat inflated. Vertex smooth, shiny, haiiless; 
vertigial sulcus rather deeply impressed, not reaching 
upper level of Ihc antennal sockets. Antennae rather 
long, slender, hardly elavate. Anienomeres 
subeylindrical; 6th slightly more obconical but not 
in flared. Pubescence moderate in proximal 
antennomeres becoming dense in distal ones. Relative 
length qf antennomeres 2 to 6: 1.00, 0.95, 0.90, 0.80, 

Collum: A little wider than head, subtrapczoidal in 
dorsal outline. Anterior border straight or very faintly 
concave, widely rounded more laterally and straight 
again towards lateral sides. Posterior border widely 
emarginate in middle, widely rounded towards lateral 
sides Lateral border moderately widely and 
symmetrically rounded. Surface transversely weakly 
convex, laterally more strongly so, and even slightly 
incurved at sides. Longitudinally surface evenly widely 
convex. Lateral margin with a narrow and low rim; 
premarainal furrow distinct, disappearing at level of the 
lateral edge of the vertex. Surface smooth, shiny and 

Somites: Rather weakly constricted. Prosornites dulled 
by a fine cellular structure and with fine striae. Waist 
narrow, rather distinctly demarcated from pro- and 
ruetasomites, dorsally finely but distinctly longitudinally 
ribbed down to the level of the paranoia, faintly striolale 
below that level. Vleratergrtes smooth or with some line 
wrinkles, shiny and hairless. Transverse furrow present 
on 5th to 16th somites, vaguely also on 17th somite. 
Purrow moderately impressed and with some vague 
sculpturing, running taterad to a distance from upper 
margin of paranoia equal to the dorso-ventral diameter 

of a poriferous paranotum. Sides smooth, or somewhat 
irregularly and finely wrinkled, shiny Anteriot somites 
up lo 4th with sides a little subgranularly uneven. No 
pleural keels. 

Paranoia: 2nd somite a little w ider than the collum. 
Laicial border in dorsal outline straight anteriorly and 
a little diverging in caudal direction, and widely convex 
posteriorly. Latero-posterior edge narrowly rounded, 
incurved so as to be slightly produced and projecting 
a little behind margin of somite Posterior border almost 
obsolete Anterior border moderately widely rounded, 
latero-anterior edge narrowly rounded, with a small, 
almost obsolete lateral tooth In lateral aspect uppei 
margin widely and weakly concave dorsally, situated on 
a low level, and therefore scarcely visible from above; 
upper margin slightly sloping in anterior direction. 
Paranotum dorsoventrally rather narrow, The upper 
furrow disrinci along all margins including the antcnor 
and posterior. Third somite a little narrower than 2nd 
and a little wider than 4ih. Paranoia in dorsal aspect 
evenly rounded, anteriorly and posteriorly more 
narrowly. Posterior edges particularly in 3rd somite 
angular, in 3rd somile slightly produced caudad. bin not 
projecting behind margin of the somite; in 4th naoowly 
rounded and not produced. Posterior border in 3rd 
somite very short, almost obsolete, in 4th obsolete. In 
lateral ysped paranoia of 3rd and 4ih somites with a 
widely concave upper margin, rather abruptly curving 
dorsad anteriorly, posteriorly curving upwards and 
shortly paralleling posterior margin of somite Dorso- 
ventral width raiher narrow 10 moderate. Lower 
demarcation formed by a depression reaching cephalad 
to about two fifths of length of metasomite. Posterior 
edges of paranoia in lateral aspect acuminate. Paranota 
of 5th and subsequent somites rather weakly prominent. 
In dorsal aspect lateral margin of poriferous paranota 
widely rounded, with a fainl emargination from pore 
area to caudal edge. Lateral rounding of poreless 
paranota a little wider. Posterior edge of paranota 
narrowly rounded, but in poreless somites and in 
poriferous somites oi caudal half of body posterior 
edges become more angular and quite weakly produced 
caudad, though not projecting behind the margin 
Pusterior border quite short, convex in poriferous to a 
little emarginate in poreless somites. Jn lateral aspect 
upper margin is faintly concave anteriorly and a little 
convex posteriorly in poriferous somites, straight or even 
faintly concave m poreless somites. Dorsal furrow 
curving a little upward anteriorly, nor reaching waist but 
running; cephalad to about four-fifths length of 
metasomite. Dorsal furrow eaudally curving abruptly 
dorsad and briefly parallelling posterior margin o\' 
somile Ventral impression demarcating the paranoia 
visible up to about two-fifths of length of metasomite 
in both poriferous and poreless somites. Dorso-vcmral 
width of paranota rather narrow, the poreless only a 
liule narrower than the poriferous. Paranoia not callous 
Pores of modcrare size, situated at anterior end oi an 


oval excavation. Posterior edges of paranoia in lateral 
aspect more or less acuminate. 

Siermies and legs: Sterniies of middle somites longer 
than wide (ratio; 1,6:1.0). Cross-impressions strongly 
developed, with deep longitudinal and transverse 
impressions. Ai bases of coxae rather large rounded 
conical protuberances projecting downward and a lirtle 
caudad, distinct particularly m anterior pair of coxae 
and especially in 9th, 10th and 11th somites, becoming 
less distinct in posterior somites. Pubescence of stemites 
dense, wilh setae of moderate length. Sternite of 4th 
somite rather widely and moderarely deeply excavate, 
moderareJy seiil'erous, with long setae. Sternite of 5th 
somite with a low hump between anterior legs, which 
is produced into a large shovel-like process a little in 
front of coxa! bases. This process is directed cephalad 
at base, crooked halfway so as to become directed more 
venrrad distally. Process projecting distinctly in front of 
sternite. Posterior surface in lateral aspect convex at base, 
more distally concave; anterior sutface convex in profile, 
apically provided wilh a dense brush of short setae. 
Process a little broader than distance between anterior 
coxae, distally quite broadly rounded, subangular in the 
middlc. Posterior surface rather weakly pubescent with 
long setae. Transverse impression distinct and deep. 
Posterior part of sternite raised as a transverse, rounded 
wall, without longitudinal impression, densely set with 
long setae. Sternite of 6th somite not raised above ventral 
level of mctasomal ring, deeply excavate. Coxal bases 
scarcely raised. Pubescence moderate, with long hairs. 
Transverse impression weak. Sternite of 7th somite with 
a large gonopod aperture pressing the ambulatory legs 
a little laterad. Latero-cephalad of aperture a rather low 
callous wall. Sternite of 8th somite anteriorly widely 
concave and only slightly raised above ventral Jevel or 
mctasomal ring. Anterior coxal bases not raised, 
distinctly more separated than posterior. Transverse 
impression weak. Pubescence dense, with long setae. 
Legs of moderate length, rather stout. Prefemur dorsally 
moderately convex, femora faintly arched. Pubescence 
dense on ventral side of all podomeres and on all sides 
of the tarsi and the anterior tibiae, remaining surlaces 
without apparent pubescence. Hairs rather long. 
Seopulae present on tibiae and tarsi of all legs up to 
6i h somite, absent on all postgonopodal legs. Relative 
length of podomeres 2 to 6 in middle somites: 0.60, 1.00, 
0.65, 0.55, 0.75, Pirst leg strongly inerassatc, with the 
usual ventral femoral tubercle. Coxae of 2nd legs ventro- 
apically thickly rounded, but not projecting. 

Anal somite: Upper profile faintly convex, almost 
straight Surface somewhat wrinkled. Epiproct of 
moderate length and thickness, slightly concave on 
ventral side* broad at base with sides concavely 
converging, more distally straight and convex towards 
apex. Near apes a slight stepwise narrowing. Apex 
truncate and a little ernarginatc, lateral edges narrowly 
rounded. Setae not on tubercles. Valves with rather 

narrow and moderately high rims. Setae on minute 
tubercles. Hypoproct large and triangular, parabolically 
rounded with sides widely and apex more narrowly 
rounded. Serae on small tubercles which do not project 
outside margin. 

I IG. 3. Someihus grosn n. sp., holotype*. right gonopod. medial 

Gonopocls: (Fig, 3) Coxa long and stout, tapering a 
little towards apex and curving a little caudad. At medio- 
anterior side of apex a small conical process. Prefemur 
rather short, rounded, its longitudinal axis transverse 
to axis of acropodite. Demarcation from aeropodite 
transverse. Acropodite split into I wo main branches: a 
caudal tibiotarsus, and a combined solenomerite and 
femoral process, Tibiotarsus long, narrow at its base bu? 
widening halfway to a laminate apical part. Combined 
solenomerite and femoral process split at about three 
quarters of length into a solenomerite proper and 
separate femoral process. Latter tapers apically and 
curves caudad. Spermal channel runs along anterior side 
of acropodite, and finally along medial side of 


KU S. At SI. ML !-. |y {}): KM? 

\hi}\ / { js:< 

solenomerite proper, making a loop in a triangular 
preapical process. Apex of solenomerite narrow, point- 
ing caudad. 

Female- Differing from the male, aside from the usual 
sexual characters, in having vertex of head transversely 
widely and evenly convex. VertigiaJ sulcus quite deeply 
impressed. Antennae shorter, with the antennomeres 
more obconical. Relative length of antennomeres 2 to 
6: 1,00, 0.95, 0.95, 0.95 t 0.90. Third somite with a low- 
transverse pleural keel along posterior margin. Siernites 
flattened, impressions distinctly less deep than in the 
male, longitudinal ones very wide. Cones absent. Ratio 
of length/width: l.hl.O. Pubescence rather dense, hairs 
shorter than in male. Legs with straight femora. Relative 
length of podomeres 2 to 6 in middle somites: 0.80, 1.00, 
0.55, 0.55, 0.70. Coxa of 2nd pair of legs distally 
produced into a large caudal transverse, callous process, 
giving coxa from behind a broad irtangular aspect, as 
broad at base as long medially, and basally produced 
laterad of pre Femoral basis. Epigynal structure 
consisting of two paramedian rather narrow 
emarginations embracing coxal bases of 2nd pair of legs. 
and medially separated by a triangular prominence. 


The new species is easily distinguished from thclype- 
species, & fttsapes* by the gonopods having a long, 
distally widening tibiotarsus (Chamberlin (1920) 
describes the "basal spur" as a 4, thin lanceolate blade 
narrowed at each end, a little twisted . . V). the absence 
of pleural keels in the male, the presence of sternal cones 
in the male, and the larger size (in & fusdpes 2.8 mm). 

The species is named after its collector, apparently 
one of the very few people in South Australia who took 
the trouble to collect millipedes, including most of the 
material treated in this paper. 

Onatcludosoma o. gen. 

Generic diagnosis 

Rather robust to small Australiosomatini with 20 
somites and a normal pore formula. Head with vertex 
transversely flattened in male, normally convex in 
felnale. Antennae of moderate length, slightly to 
distinctly elavate, basal antennomeres subcylindrical, 
distal ones more obconical in shape. 

Somites rather weakly to moderately constricted, 
waist rather narrow* distinctly longitudinally ribbed or 
beaded. Metatergitc.s smooth, hairless. Transverse tin row 
present from 5th somite onwards, rather weakly to rather 
deeply impressed. Pleural keels vestigial or absent in 
mule, moderately developed up to 4th somite in female. 

Paranoia rather weakly developed. 

Sternites distinctly longer than wide in male, about 
as long as wide or a little longer than wide in Female. 
Sternal cones weakly developed to vestigial. Sternite of 
5th somite of male with a process between anterior legs. 

Legs tather long; first leg of male incrassate and with 
a ventral femoral tubercle, Tibial and tarsal seopulae 
present only in a number of legs in the anterior half 
ol the body of male. 

Gonopods with pretcmur ovoid, its longitudinal axis 
almost transverse on the axis of the acropodhe. 
AcTopodile deeply split into two main branches, femora] 
pail almost vestigial. Tibiotarsus undivided, more or less 
clubshaped, with a rather narrow "stern" and a widened, 
more or less complicated distal half. Seminiferous 
branch exceeding tibiotarsus in length, end typically 
uncatc and recurved. Spermal channel running along 
posterior side ol seminiferous branch, turning distallv 
towards medial side, and running to extreme distal part 
of the seminiferous branch before abruptly recurving 
towards apex. Just proximad of distal edge of 
seminiferous branch and ccphalad of course of spermal 
channel there is a vestige of the femoral process, which 
is completely absent, however, in the type-species. A little 
more proximad on mediocaudal side of seminiferous 
branch there is in the type-species a short spioelike 
process, vestigial or absent in the other species, 

lype-spedes: Atisfrafiosoma casnmeum Attems, 1944. 


This genus is characterized mainly by the peculiar 
shape of the aciopodite of the gonopods, with its single 
clubshaped tibiotarsus, its typically uncatc seminiferous 
branch, the absence or almost complete reduction of 
the femoral process, and the course of the spermal 
channel, which makes a slight loop before running 
towards the apex of the solenomerite. 

In having the gonopod telopodite deeply split into two 
mam branches, Oticocladosoma bears some similarity 
to the genera Didadosowa Brolemann, 1913, 
Dtdadosomdla Jeekel, 1982, Phyllodadosoma Jeekel, 
]%8, and Somethus Chamberlin, 1920. Dicladosowa, 
from Mt. Kosciusko, is distinct in having the base of 
the tibiotarsus twisred towards the laiero-antenor side 
of the seminiferous butneh. It moreover seems distinct 
in having apparently no loop in the course of the 
spermal channel. Dtdadosomelfa, Phyllodadosoma and 
Somethus have a more erect telopodite, and the spermal 
channel makes a loop in a distinctly developed process 
Dicladosomelfa, from southeastern New South Wales, 
hvi:- a more laminate tibiotarsus, and the apex of the 
solenomerite is typically directed distad 
Phyllodadosoma, from northeastern New South Wales 
and southeastern Queensland, resembles Dtdadosomdla 
in mosl of the characters mentioned, but it is 
distinguished by the abbreviate solenomerite, ihe apex 
of which extends scarcely beyond the loop-bearing 
process, Soffiethas is distinguished by having a distinctly 
developed femoral process, Both, Didadosomdla and 
Phyllodadosoma, lack any indication of a fcmoial 

The new genus eoniains, besides its type-species, 
which is divided into two subspecies, two new species. 

Mil I |F»fcW£S !k()M AUSIkM IA 5 


Geographically it seems to be confined to the Ml. Lofty 

Oncodadosoma castaneum (Attems) 

Australiosoma castaneum Attcms, 1944; 249; Jeekel, 
1968- 26. 

This species is represented by two subspecies, O, c. 
castaneum and O. c. ingens n subsp., characterized by 
a significant difference in size 


Q c. custaneum (Attems): 

ML Lofty, 2i.iX.1883, leg. Teppei; I S , i v 

Belair. I5V.1938, leg. R. V Southcott, ZS , 

Belair, Mt. Lofty Ranges, leg. N. B. Tindaie, Id" . 

Q c. ingens n. subsp.: 

Near Mt. Lofty Station, 1V.I883, Dr. Haacke don. )C f 

holotype, 23 £ , 11? ,6 juvc? (19 somites), 2 juv. i 

(19 somites) paratypes. 

Bridgewater, 2.IL1884, leg. Teppet, 1 <1 paratype. 

Norton Sumrnit> Mt. Lofty Ranges, 7.1V.1884, leg. 

Tepper, 2 cf paratypes. 

Mt. Lofty. Waterfall Gully, 23.VI 1884. leg, Tepper, I cf 


Onkapannga River, near Mylor, 1I.V.1947, leg, G P 

Gross, 2^ paratypes. 

Mylor, 20.1YI957, leg. G. F. Gross, 3 o* paratype. 

Upper Stun, Soil T\ 104, 5.V1962, leg. R. V. Southeort, 

I & paratype. 

Bridgewater, Lot no. 1l-72a, LX.1972, leg. G. H. Baker, 

I <J , J 9 paratypes. 

Without locality or other data, 9^ , 5 % , 1 juv i (19 

somites) paratypes. 

Without locality or other data, 2^,2 ? paratypes. 


Co/our; Head brown, vertex and lateral scienter 
blackish. Area around the antennaJ sockets and margins 
of lateral sclerites pale brownish. Antennae brown, 6th 
antennomere and basal part of 7th infuscate; tip whitish; 
the intersegmental membranes pale brownish. Collum 
black isfu margin behind vertex reddish brown; a rather 
narrow /one along posterior margin, broadest medially 
and tapering lowards sides, brownish yellow. Prosoniites 
and anterior part of metasomites blackish brown, 
posterior part of metatcrgitcs, i.e. medially from just in 
Iront of transverse furrow eaudad, yellowish brown. 
Posterior part of paranota and posterior zone of sides 
also yellowish brown. Demarcation between the darker 
and lighter colours not sharp. Venter, sternites and legs 
brownish, three distal podomeres infuscate, inter- 
segmental membranes and tip of tarsi yellowish brown. 
Anal somite dorsally, including epiproct, yellowish, sides 
blackish. Paraprocts black, margined with yellowish 
brown. Hypoproet yellowish brown. More heavily 
tnfuscated specimens have pale colour only along 
posterior margin of metasomites, Specimens which have 

been stored in alcohol for a long time are brown in 
general, with the lighter colour only vaguely indicated. 
Juveniles dirty whitish, without colour pattern. 

Width: O. c. castaneum; J : 2,8-3,2 mm, * : 3,0 mm. 
Q C ingens: c <: 3,5-4.3 mm, £ : 3.4-4.0 mm, juv c? (19 
s.): 2.9-3.2 mrn, juv * (19 s.); 2.7-2.9 mm. 

Head and antennae; Labrurn moderately widely and 
moderately deeply ernarginate. Clypeus strongly 
impressed towards labrurn, surface uneven due to 
presence of setiferous pits; on each side below antenna! 
sockets wide impression. Lateral border of clypeus 
faintly convex, with a much near labium. Pubescence 
moderate becoming sparse in frontal region and on 
lateral sclerites; above labrurn a series of hairs, hairs 
rather short. Frons not demarcated from vertex or 
clypeus. Antennal sockets separated by L4 times 
diameter of a socket or by 0.7 times length of 2nd 
antennomere. Vertex shiny, hairless, transversely faintly 
convex, a little mote so near lateral edges, longitudi- 
nally almost evenly and rather widely convex. 
Postantennal groove rather deep and rather wide; wall 
moderately prominent. Beanshaped area at posterior 
side of antennal sockets well demarcated and inflated. 
Vertigial sulcus well impressed, not reaching upper level 
of antennal sockets, with short fine transverse wrinkles. 
Antennae of moderate length, moderately stout, 
slightly clavate, Antennomeres subcylindrieul, but 5th 
more obconical, and 6th still more obcomcal but not 
inflated. Pubescence moderate in proximal anten- 
nomeres. becoming dense in distal ones. Relative length 
ol 2nd to 6th antennomeres: 0.9?, LOO, 0.95, 0.85. 0.70. 

Collum: Subtrapezoidal in dorsal outline, a little wider 
than head. Anterior border straight in middle, widely 
rounded more laterally and straight again towards lateral 
sides. Posterior border widely and weakly concave in 
middle, straight laterally. Lateral sides asymmetrically 
and rather widely rounded, with strongest convexity 
caudal Surface hairless, shiny and almost polished, with 
weak irregular wrinkles. Surface transversely widely 
convex, more strongly so towards lateral sides and 
laterally even slightly incurved; longitudinally weakly 
convex, slightly more so towards anterior and posterior 
margins, Marginal rim laterally moderately wide, not 
particularly raised; premarginal furrow distinct, 
disappearing at lateral edge of vertex. 

Somites; Constriction rather weak. Waist rather 
narrow, sharply demarcated from pro- and metasomites, 
distinctly longitudinally ribbed down to upper level of 
paranota, finely striate below that level. Prosomites dull, 
with a fine cellular structure and fine short striae. 
Metatergites smooth, hairless, shiny, with some irregular 
weak wrinkles. Transverse furrow finely and not deeply 
impressed, without apparent sculpturing, present from 
5th to 17th somite, vaguely indicated on 18th, Furrow 
running laterad to about a distance from upper 
demarcation of paranota of about one and a half times 


kt i . S\ AIM. Ml IN [<J |?J V) 1? 

A A/ 1. tftfj 

or a little more the dorsovcntral width of a poriferous 
paranotum. Sides smooth or slightly wrinkled, granulate 
up to 4th somite. Pleural keels abortive or absent. 

Panmotu: 2nd .somite a little wider than collum; 3rd 
somilc a little wider than 2nd and about as wide as 4th. 
Paranoia of 2nd somite with anterior margin a little 
thrust forward, widely convex. Latero-antcrior edge 
subangular, narrowly rounded, without distinct tooth. 
Lateral margin widely and almost evenly rounded, but 
behind middle a Hide more strongly convex arid eaudally 
straight, hardly diverging. Latero-posterior edge 
subangular, slightly produced and projecting slightly 
behind posterior margin of somite. Posterior margin 
short, widely convex. Paranoia situated on a low level. 
rather weakly prominent, although margin visible from 
above. Upper margin in lateral aspect widely and weakly 
convex, sloping a little cephaiad. Marginal callus rather 
wide from the middle onwards, narrower anretiorly, its 
lower side straight. Premarginal furrow distinct on all 
sides, briefly parallelling the posterior margin of the 
somite. Paranoia of 3rd somite with anterior margin not 
shouldered at base, widely convex, merging via a 
stronger rounding into the widely convex lateral margin. 
Posterior edge subangular, slightly produced caudad. 
Posterior margin short, a little concave In lateral aspect 
marginal callus moderately wide; ventral demarcation 
by a depression is present only in posterior half, 
converging dorsad somewhat eonvexly and meeting 
dorsal demarcation in an acute angle. Paranota ot 4th 
somite similar to those of 3rd, but lateral margin a little 
more widely rounded, and posterior edge eaudally not 
produced. Paranota of 5th and subsequent somites 
rather weakly prominent, Lateral margin in dorsal aspect 
weakly to faintly convex, anteriorly slightly more 
rounded; poriferous paranota eaudally slightly more 
convex than the poreless. Latero-posterior edge 
subangular, quite narrowly rounded, faintly produced 
caudad in most somites, more so in the somites of 
posterior half of the body, but projecting behind the 
caudal margin of the somite only in the 18th. In lateral 
aspect upper demarcation of the callus convex 
(poriferous somites) or straight (poreless somites), 
premarginal furrow turning upward near waist but not 
reaching it. Vennat demarcation by a depression present 
only in the caudal half or three- fifths of the paranoia, 
converging straight (poreless) or convexly (poriferous 
somites) towards the dorsal demarcation in an acute- 
angle, more acute in poreless than in poriferous somites. 
Dorsovcntral width of paranota moderate, in poreless 
somites about two-thirds of poriferous Posterior margin 
finely rimmed, the premarginal furrow briefly 
parallelling caudal margin of metatergitc. Pores situated 
quite near the ventral edge of the marginal callus, in 
a rather small oval pit. 

Siertvtes and legs: Stcrnitcs of middle somites longer 
than wide (ratio J.6:l 0). Cross impressions well 
developed, rather wide. Sternal cones present on all 

postgonopodial stemites up to 17th somite, rathei small 
near anterior coxae, and still more weakly developed 
near posterior coxae. Pubescence rather dense to dense. 
located in four quadrants of each stci mte Median and 
transverse impressions hairless; hairs of moderate length. 
Sternile of 4th somite rather broad, with a well- 
developed median impression, no sternal cones. 
Pubescence rather dense. Sternite of 5th somite with a 
subtrape/.oidal process arising between and a liltlc in 
front of anterior coxal sockets; process broader than 
long, a little wider than width between coxae, directed 
downward and cephaiad, bending abruptly downward 
in distal hall, projecting distinctly in front of anterior 
margin of stemilc. Anterior surface of process- in profile 
convex hallway, apex with a dense brush of short setae. 
Posterior surface widely concave, moderately densely set 
with rather long setae. Apex of process in caudal aspect 
broadly rounded, medially faintly angulatc, lateral edges 
narrowly rounded. Behind process a deep transverse 
impression. No longitudinal impression between 
posterior coxae, but posterior coxal sockets connected 
by a transverse wall which is densely set with long setae. 
Sternite of 6th somite scarcely (anteriorly) or not 
(posteriorly) raised above ventral levei of mctasornal 
ring. Coxa! sockets scarcely raised and coxae not 
elongate. Pubescence consisting of two transverse zones 
of densely set long hairs. Sternite of 7th somite 
lateroanteriorly with a finely rugulose-granulose 
transverse wall. Gonopod aperture large, separating 
coval sockets of posterior legs, Stctntte of 8th somite 
anteriorly not raised above ventral level of metasomal 
riijg; anterioi cosac somewhat separated. Transverse 
impression weakly developed. Posterior part not 
modified, cones abortive. Pubescence moderate, hairs 
rather long. li>gs in general rather long and stout, 
prefemora dorsally convex, femora faintly arched. 
Pubescence ventrally dense on ail podomcres, hairs 
rather long. Dorsal pubescence not obvious, except on 
tibiae and tat si and partly also on posifemora. Scopulac 
of tibiae and tarsi present up to anterior k'gs of 8th 
somite, totally absent from posterior legs of t hat somite 
onwards. Relative length of podomeres 2 to 6; 0.80, 1.00, 
0.65, 0.60, 0.80. Legs of first pair strongly incrassate, 
with a ventral femoral process. Coxae of 2nd pair 
medially rather Thickly rounded but only weakly 
produced distad. 

Anal somite: Upper profile about straight or faintly 
convex. Epiproct broad, moderately thick, rather short, 
ventrally somewhat concave. Sides concavety converging, 
narrowing abruptly just before apex, which is truncate, 
eaudally weakly emarginate, lateral edges narrowly 
rounded. Setae not on tubercles. Paraprocts with narrow 
and rather low rims; setae not on tubercles- or latter 
abortive. Hypoproct parabolically rounded, setae on 
abortive tubercles. 

Gonopods: (Figs 4-5) Characterized mainly by the 
shape of the tibiotavstis. Moreover there is no trace of 



FIGS. 4-5. Oncududtisoma tmtancwn ingenS n. subsp., holotypeo* 4: nghi tionopod, medial aspect. 5: telopodke of left gonopod. anterior aspect, 

a femoral process. On the other hand the species has 
a spinelike process arising from ihe medio-caudal side 
of the seminiferous branch a little proximad of the base 
of the distal hook. The morphological status of this 
process is not yet clarified. 

Female; Head with the antennal sockets separated by 
1.4 times diameter of a socket or by 0.8 times length 
of 2nd antennomere. Vertigial sulcus slightly less 
impressed, vertex transversely widely and evenly convex. 
Relative length of antennomeres 2 to 6: 1.00, 0.95, 0.90, 
0.85, 0.80. Somite 2 with pleural keels represented by 
a strongly developed rounded ridge. A thick dorsally 
sharply demarcated rounded swelling in 3rd somite, in 
4th an abortive swelling. Sternites about as wide as long. 
The cross-impressions much weaker, without sternal 
cones. Setation rather dense, but setae rather short. Legs 
with ventral pubescence rather dense, hairs of moderate 
length. Relative length of podomcres 2 to 6: 0.70, 1.00, 

0.50, 0.45, 0.80. Coxa of 2nd pair of legs with a caudo- 
lateral callous process, pointed and directed caudo- 
laterad, medially not produced. Epigynal structure 
consisting only of two emarginations of ventral side of 
3rd somite, embracing coxae of 2nd pair of legs, and 
medially separated by a low triangular process pointing 
cephalad. Lateral border of emarginations raised. 


When we compare the present drawings of the 
gonopods with the one which Attems (1944) published, 
there can be but little doubt that the presently recorded 
material belongs to the same species. It will be noted, 
however, that there are important discrepancies between 
the drawings. In the first place Attems misidentified the 
small process at the medio-caudal side of the 
seminiferous branch as a side branch of the tibiotarsus 
(the latter erroneously called "Fcmoralfortsatz"). 


KM. S. AUM, MI'S. 1911); 19-37 

Muv. tm 

Furthermore, he illustrated a separate femoral process 
(erroneously indicated a.s 'Tibiotarsus"), emanating 
distinctly proximad ol the distal edge of the seminiferous 
branch. Such a process is not observed in the present 
material, and we must assume that Auems either 
misinterpreted thechitinous structures in his preparation 
or that the gonopod he examined was damaged. 

Tiie subspecies ingens appears to differ from the 
nominate subspecies mainly in its larger size. Differences 
in the details of the gonopods must be substantiated 
by future research when more material becomes 
available, Whereas the subspecies castaneum appears to 
be confined to Belair (unfortunately Auems gave only 
"Sudaustralietf as type locality), the subspecies ingens 
occurs in a much larger area of the Mr Lofty Ranges. 

Oncocladosoma conigerum n sp, 


Without locality label, but found in a tube together with 

O. castaneum ingens: <? holotype, 4 £ paratypes. 

Near Mi. Lofty Station, I V.IS83, Dr Haacke don., I e 


Without locality label, 6 <s paratypes, 2 ? paratypes. 


Colour: Essentially the same as that of the preceding 
species, but less infuscate (possibly duo to prolonged 
preservation in alcohol). Dark colour of head confined 
to vertex and frontal region; lateral sclentes of the head 
pale brownish. Antennae brown, joints paler Colktm 
and somites as in castaneum but pale and dark colours 
less contrasting. Distal podomeres scarcely infuscate. 

Width <? : 2.1-2,4 mm; S : 2.3-2.6 mm. 

Head and antennae: Lateral border of clypeus with 
notch indistinct, Antennal sockets separated by 1.3 times 
diameter of a socket or by 0.55 times length of 2nd 
antennomere. Postantennal bean-shaped area a little 
inflated, moderately demarcated. Antennae rather stout. 
Relative length of antennomeres 2 to 6: 1.00, 0.95, 0.80, 
0,80, 0.75. 

Collum: A little narrower than head. Lateral sides 
almost evenly rounded. 

Somites: Waist distinctly beaded dorsally. Transverse 
furrow disappearing laterally at a distance from dorsal 
demarcation of paranota equal to dorso-ventral diameter 
of 41 poriferous paranotum. Pleural keels weakly 
developed up to 3rd somite, faintly indicated in 4th. 

Paranota: 3rd somite about as wide as 2nd; 4th a little 
wider than 3rd. Paranota of 2nd somite with marginal 
callus of equal width. Paranota of 3rd somite without 
anterior edge, latero-anterior border rather strongly 
rounded. In lateral aspect upper demarcation of 
paranota of 3rd and 4th somites slightly concave, turning 
upward anteriorly and posteriorly; the upper 
demarcation not sloping caudad. Paranota of 5th and 
subsequent somites with posterior edge narrowly 

rounded, scarcely produced except very shghrly so in 
posterior somites but not projecting behind posterior 
margin of somites. In lateral aspect ventral demarcation 
of paranoia more strongly convex, meeting upper 
demarcation in a wider angle than in castaneum, giving 
posterior edge of paranota a more subiruncate 

Stermtes and legs: Sternites oi' middle somites longer 
than wide (ratio 1.65: 1.00). Process of sternitc of 5th 
somite with process about as long as wide at base, about 
parabolical in posterior aspect. Posterior half of sternite 
with a median rather deep furrow; pubescence rather 
dense, setae longish. Sternite of 6th somite with a 
distinct transverse furrow, but without longitudinal 
impression. Sternite o\ 7th somite with a weak callus 
latere- eephalad of gonopod aperture. Legs with 
scopulae up to first pair of 8th somite as in preceding 
species. Pubescence on ventral side of legs long. Relative 
length of podomeres 2 to 6 in middle pan of body: 0.65, 
1.00, 0.60, 0.55, 0.80, 

Anal somite: Upper profile faintly convex, weakly 
convex in front of base of epiproct. Fpiproct longish, 
broad. Caudal emarginalion very weak. Hypoproct 
raihcr large, about evenly .scmicireularly rounded. 

Gonopods: (Figs. 6-7) Coxa relatively a little more 
robust than in O. castaneum. Prefemur with a weakly 
chitinized conical process emanating from 
intersegmental membrane connecting coxa and 
prefemur. Acropodite largely similar to (hat of O. 
castaneum. differing mainly in shape of ttbiotarsus. 
Seminiferous branch with a slight vestige of a femoral 
process, The larger spinelike process of O. castaneum 
is represented here by a tiny tooth 

Female: Head with clypeus moderately impressed 
towards labrum. Antennal sockets separated by I 3 times 
diameter of a socket or by 0.7 times length of 2nd 
antennomere. Antennae relatively shorter; relative length 
of antennomeres 2 to 6: LOO, 0.90. 0.85, 0.90, 0.75. 
Collum more evenly rounded transversely. Somites with 
pleural keels on 3rd somite represented by a low cone 
near caudal margin. On 4th somite a faint ridge- 
Sternhes of middle somites 12 limes longer than wide. 
Legs less incrassate, femora straight. Relatfve length of 
podomeres 2 to 6 in middle part of body: 0.70, LOO, 
0.55. 0-50, 0.70. Epigynal structure with paramedian 
cmarginations deeper than in castaneum surface of the 
ventral side of 3rd somite anteriorly raised, and median 
conical process directed more downward. 


In the characters not mentioned above, the description 
of O. castaneum applies. 

This species looks like a diminutive form of O. 
castaneum, being even smaller than the subaduh 
specimens of that species, It differs only in small 
proportional details of its external morphology, but it 
is well characterized by the structure of the gonopods. 



FIGS. 6 7. Oncockuhsoma comgerum n. sp., holotyped-. 6; righi gonopocl, medial aspect, 7: left gonopod, anterior aspect 

Although most of the material was lacking a locality 
label, the fact that part of it was found in a tube 
containing O. castaneum ingens seems to imply that it 
occurs sympatrically with that species in the Mt. Lofty 

Oncocladosoma clavigerum n. sp. 

Belair, VI. 1883, leg. J. W. Haacke, <f holotype, \5£ , 

8f , 2 juv. & (19 somites), 1 juv. ¥ (19 somites) 

pa rat y pes. 

Blackwood, VI. 1883, leg. I W. Haacke, 2d', 1 <? 


Woodley's Vineyard (near Glen Osmond, SE Adelaide), 

soil, Berlese funnel, 30V1I-8.V11I.1950, leg. R. V. 

Southcott, 2 cf paratypes. 

Sellick's Hill, 23.IX.1954 (E.S.I. 1592), leg. G. F. Gross, 

1 d\ 


Colour: Similar to that of the two preceding species, 
but less pronounced possibly due to preservation. Dark 
colour of head confined to vertex and frontal regions; 
lateral sclerites of head pale brownish. Antennae dark 
brown, distal annul i of antennomeres and 
intersegmental membranes pale brownish. Collum 
brown, without distinctly paler marginal areas. Somites 
brown, darkest in waist area, without paler bands; only 
posterior halves of paranoia a somewhat paler brown, 
and posterior halves of metatergites also slightly paler. 
Anal somite also with a paler epiproct, and margins 
vaguely paler. Hypoproct pale brown. Sternites and legs 
pale brown; three distal podomeres somewhat infuscate, 
but no pale annuli. Tip of tarsi pale. 

Width: & : 1.5-1.8 mm,? : 2.0-2.3 mm, juv. rf (19 
somites) 1.4 mm, juv. 9 (19 somites) 1.4 mm. The 


KL(_ \ MiKI MIS. \$m IMS 

W<n ««.1 

material from Belair seems slightly smaller (<J ; 1,5-1,8 
mm, v . 2.0-2.2 mm) than the specimens from elsewhere, 
which measure 1.8 mm and 2,3 mm for the two sexes 

Ifeod and antennae: Labrum rather widely 
emarginate. Clypeus rather strongly impressed towards 
labrum. Antenna I sockets separated by 1.6 limes 
diameter of a socket or by 0.85 times length oi 2nd 
antennomere. Antennae rather stout, distinctly clavate 
(involving 5th and especially 6th antennomere), 6th 
antennomere a tittle inflated Kelative length of 
•anrennornercs 2 to 6; 1.00, 1.00, 0.95, 0.95, 0.90. 

Cotlum; Distinctly narrower than head, subreniform 
to subtrape^oidal in dorsal outline. Anterior border 
almost evenly rounded, only slightly more convex behind 
edges of vertex. Posterior border laterally widely convex. 
Lateral sides rather narrowly and almost evenly rounded. 
Surface of cullum longitudinally evenly and widely 

Somites; Constriction moderate. Waist dorsalJy 
distinctly beaded, laterally distinctly to finely striate. 
Prosomitcs dull, with rather pronounced fine cellular 
structure. Transverse furrow of mctatergites sharply and 
rather deeply impressed, with fine longitudinal striatioru 
disappearing laterally at about a distance from dorsal 
demarcation of paranota equal to dorso-venttal diameter 
of a porejess paranotum. Transverse furrow indicated 
on 4th, and very vaguely also on 18th somite. Pleural 
keels on 3rd somite weakly indicated by a furrow; 
abortive on 4th somite. 

Paranota: 2nd somite scarcely wider than collum. 
Paranoia of 2nd somite with latero-antcrior edge 
narrowly rounded, without tooth. Lateral margin only 
faintly convex, a little more so towards posterior edge, 
scarcely visible from above. In lateral aspect margin 
almost straight, moderately thick and of equal width, 
Paranota of 3rd somite laiero-anteriorly evenly and 
rather widely convex. Posterior edge angular, slightly 
produced caudad bui not projecting behind margin of 
the somite. Paranota of 4th somite with posterior edge 
faintly produced. In lateral aspect dorsal demarcation 
of paranoia of 3rd and 4th somites faintly concave, 
turning upward gradually at anterior end. Paranota of 
5th and subsequent somites with ventral demarcation 
rather convcxly converging with upper demarcation, 
giving paranota, particularly the poriferous ones, an 
obliquely truncate aspect in lateral view. Cauda! edges 
not projecting behind margin of somites. Pores in a 
relatively large and deep pit, nearer ventral margin, 
rather near posterior edge. 

Sternites and legs: Sternites of middle somites 
longer than wide (ratio 1.6: 1.0). Sternal cones not 
obvious, quite weakly developed Pubescence moderately 
dense, all over sterniie, the setae of moderate length. 
Sternite of 5th somite with the process between the 
anterior legs subtriangular, the apex rather widely 

rounded, about as long as wide. Anterior side in profile 
weakly convex, the process almost perpendicular, 
scarcely projecting in front of sternite, Caudal half of 
sternite widely transversely concave, rather densely set 
with longish setae. Sternite of 6th somite with coxal 
sockets conspicuously more widely separated. Stemite 
of 7th somite with pregonopodial wall weakly 
prominent. Sternite of 8th somite with anterior coxae 
widely separated. Sternite anteriorly flattened, a little 
raised. Transverse impression and posterior half of 
stern iic not modified. Pubescence moderate, with setae 
longish. Legs oi' moderate length or longish. rather 
stout. Relative length of podomeres 2 to 6 in middle 
somites: 0.75, LOO. 0.65, 0.70. 0.85. Coxae of 2nd pair 
not medially produced. Coxae of first pair of legs of 
6th somite and ol anterior pair of 8th somite slightly 
produced inio a weak rounded cone. Ventral pubescence 
of legs rather dense 

Anal somite: Upper profile straight, slightly convex 
anteriorly. Apex of cpipract relatively narrowly truncate, 
scarcely ernarginatc. Hypoproct subtrapezoidal, with 
median part of apex somewhat triangularly produced; 
setae on faint tubercles which are faintly produced. 

Gonopods: (Figs 8-9) Coxa relatively slightly more 
slender than in the other two species. Tibiotarsus 
typically club-shaped, with a relatively long stem. 
Seminiferous branch characterized by the rather strong 
curvature; femoral process reduced to a tiny lobe. The 
more proximal spine, present in O. castaneutn and 
vestigial in O. conigeruni Ls totally absent here. 

Female: Head with anicnnal sockets separated by L5 
times diameter of a socket or by 0.9 times length of 2nd 
antennomere. Relative length of antennomeres 2 to 6: 
LOO, 0.90, 0.85, 0.90, 0.85. Pleural keels of 2nd somite 
well developed, those of 3rd somite represented by a low 
conical swelling near posterior margin; in 4th somite this 
swelling is very weak and small. Keels are dorsally not 
demareared by a furrow. Sternites of middle somites LI 
times longer than wide, Setae of moderate length. 
Relative length of podomeres 2 to 6 in middle part of 
body: 0.80, LOO, 0.55, 0.45, 95 Coxae of 2nd pan of 
tegs on caudal side with a low rounded transverse callus. 
Epigynal structure with median point directed obliquely 
downward; lateral anterior projection also directed 
obliquely downward. 


In points not mentioned the description of O. 

castarteum applies. 

This is the smallest representative o\' the genus 
Oncoehtdosoma, differing from the other species in 
small details of the external morphology and 
particularly in the structure of the gonopods. 

It may be of interest to note that the specimens for 
Belair are slightly smaller on the average, parallelling 
the conditions in O. cc^tatwitm. Ln the present case, 
however, the differences seem insignificant. 



FIGS, 8-9, Oncodudosoma dimgemm n. sp., lwlotypecf . S; right gonopod, medial aspect. 9: left gonopod, anterior aspect. 


The taxonomic position pf this species, allegedly based 
on a female specimen from Adelaide, has been uncertain 
since the publication of its description. Through the 
kindness of Dr M Moritz of the Museum fur 
Naturkunde der Humboldt-Universitat in Berlin, I was 
able to re-examine the type-specimen, hoping that it 
could be associated with one of the paradoxosomatids 
received from the South Australian Museum. 

The type-specimen is in bad condition; its antennae 
and most of its legs arc missing. Moreover, it is not an 
adult female, but a juvenile with 19 somites. 
Nevertheless, on account of its external characters it has 
become clear that the species is different from other 
Paradoxosomatidae reported from South Australia. In 
case the species is rediscovered in South Australia its 
most salient characters are mentioned here. 

"Strongytosoma" innotatum (Karsch) 

Polydesnius (Stran#ylosomu) innotatus Karsch, 1SS1: 42. 
Strongylosoma innotatum; Attems, 1898: 307. 


Adelaide, Schomburgk leg., 1 juv. ? (19 somites), 
ZMB Kat.Nr. 560, holotype. 

Descriptive notes 

Colour: Dull brownish, apparently Taded. 

Width: 3.5 mm. 

Head and antennae: Without particulars. 

Collum: Wider than head (Fig. 10). Lateral sides 
flaring, rather widely rounded and without posterior 
edge; lateral border merging gradually into caudal 

Somites: Waist rather narrow, without apparent 
sculpture. Metatergitcs of 4th to penultimate somite with 
a generally deeply impressed transverse furrow, running 
far laterad and reaching in most cases dorsal 
premarginal furrow of paranota. 

Rtranota: (Figs 10-11) 2nd somite distinctly wider than 
collum; 3rd and 4th each slightly narrower than 
preceding somite. Paranota of 2nd somite on a low level, 
well developed; anterior border shouldered at base and 

RB , S. AUS1. MUS. |»{3*: 19-37 

Muv. tm 

FIGS. 10-11- "Stmngyfosonia" mnotatum (Karsch), holotype V juvenile 
(19 somites). 10: left side or head, collurn and 2nd and 3rd somites, 
dorsal aspect. 11: left side of 9th somite, dorsal aspect. 

thrust forward a little, widely rounded, merging into 
rather faintly convex lateral margin via a somewhat 
stronger rounding. Posterior edge rather narrowly 
rounded but not angular. Posterior border a little 
convex; posterior part of paranota projecting distinctly 
caudad of margin of somite. Paranota of 3rd and 4th 
somites also prominent, widely rounded anteriorly and 
laterally, more narrowly rounded caudally and project- 
ing a little behind margin of somites. Paranota of 5th 
and subsequent somites at first subangular caudally, but 
in second half of body becoming more distinctly 
angular, though produced caudally and projecting only 
a little from 15th somite onwards. 

Sternites and legs: Sternites without distinctive 
characters; legs mostly absent (in specimen). 

Anal somite: Epiproct moderately developed, apex 
truncate with a weak caudal cmargination. Hypoproct 
rather broadly triangular, with sides and apex rounded. 


Since the specimen is a juvenile, most of its external 
characters may be less pronounced than in the adult, 
in particular the adult male. The paranota are different 
from those of any of the other South Australian 
paradoxosomatid genera. In particular the relatively 
wide expansion of the paranota of the 2nd somite seems 
characteristic. In this respect the species appears to have 
a certain similarity with Otoplacosoma hivittutum 
Verhoeff, 1924, from the Kimberley district of Western 
Australia, which would mean that eventually it may 
prove to belong to the tribe Antichiropodini. It should 
be borne in mind though, that it is also possible that 
the locality label is wrong, in which case the systematic 
position of the species probably will be an enigma for 


With the completion of the study of the Para- 
doxosomatidae of the collection of the South 
Australian Museum, it may be useful to summarize the 
results which have been obtained so far. 

The number of known species has increased from two 
to ten and one subspecies. The list is as follows: 
Tribe Antichiropodini 

Antichiropus mammillifer Jeekel 

Aulacoporus pruvoti (Brolemann) 

Tridaetylogonus obscunts Jeekel 
Incertae sedis 

"Strongylosoma" innotatum (Karsch) 
Tribe Australiosomatini 

Heierocladosoma zehratum n. sp. 

Heteroctadosoma gaiaxias n. sp. 

Somethus grossi n. sp. 

Oncocladosoma c, castaneum (Attems) 

(Jncocladosoma c. ingens n. subsp. 

Oncocfadosoma conigerwn n. sp. 

Oncocladosoma clavigerum n. sp. 

Considering the fact that the southeastern part of 
South Australia, where these paradoxosomatids were 
obtained, is quite remote from the nearest occurrence 
of the family elsewhere and is moreover drier than the 
areas in eastern Australia where the majority of the 
other described species occur, it is remarkable that the 
number of endemic genera is so relatively low. To this 
category belong only Tridaetylogonus and 
Oncocladosoma. The other genera, viz. Antichiropus, 
Aulacoporus, Heierocladosoma and Somethus, are 



shared with other Australian states, where, according to 
our present knowledge they all show a greater diversity. 
Antichiropus has quite a number of species occurring 
along the west coast of Western Australia. 
Heterocladosoma has three distinct species in the coastal 
area of Queensland from Cairns to Brisbane, and, 
according to as yet unpublished data, Somethus has two 
species in Tasmania and one in eastern Victoria. 
Aulacoporus has a number of species in Queensland and 
the northern part of New South Wales, but the South 
Australian record of A. pruvoti probably represents an 
introduced population, it is evident anyway that the 
fauna of South Australia, as far as the non-endemic 
genera are concerned, is composed of a mixture of 
western (Antichiropus), northeastern (Heterocladosoma), 
and southeastern (Somethus) elements (Fig. 12). 
Considering the diversity within each of these genera 

elsewhere it seems probable that South Australia became 
populated along these routes, rather than the reverse. 

The relationship of the two endemic genera has not 
yet been satisfactorily determined. Tridactylogonus 
seems to belong to a group of small antichiropodine 
paradoxosomatids which focuses in Victoria, Tasmania 
and southern New South Wales. Oncocladosoma 
appears to be closely related to Somethus, 
Diciadosometla and Phyllocladosoma, indicating a 
faunistic connection between South Australia, Tasmania, 
eastern New South Wales and southern Queensland. 

Oncocladosoma represents, moreover, a particularly 
interesting case. The genus has been found only in the 
Mt. Lofty Ranges, where it is represented by four partly 
sympatric taxa. No other paradoxosomatids have been 
reported from this probably best-explored part of South 
Australia, and the faunal composition shows the aspects 

TIG. 12. Map showing the distribution of the non-endemic South Australian ausiraliosomatine genera Heterocladosoma (*), Somethus f « 
and Antichiropus (o), and the endemic genera Oncocladosoma (L) and Tridactylogonus ( ). 


1:1 i v aum. WUSj IV (1); IM7 

AM'. /M5 

of that of a long isolated island. During an earJy petiod 
this area must have been populated by a single species, 
possibly from the Sumefhus-$t?m> which, in the course 
of a long period of isolation, radiated in a number o\' 
closely related taxa. 

The present picture of the distribution of the South 
Australian Paradoxosornatidae, incomplete as it 
undoubtedly is, seems to indicate thai the fauna is the 
result of exchanges during at least two widely separated 
periods in history. The first of these periods, which may 
he more or less arbitrarily dated at the late Mcso/oic 
or early Tertiary, saw a fauna! connection between 
Victoria and southern New South Wales and South 
Australia, and is witnessed by Trkiach logonus and 
Oncvcladosotua, The second, possibly occurring during 
the late Tertiary, brought representatives of 
Heterucladosoma, Somefhus and Au/ichiropus to South 

Of course, this is a very simplified explanation of 
what actually may have happened, but it will be 
interesting to assemble further data to verify and work 
out the details o[ the hypothesis. 

Burnside, 12.VI.1962, leg. Mr Waxman, 26, 3?. 
Burnside, 24. U 963. leg. Mr Waxman. 1c?, 16 v. 


This species was originally described from Portugal, 
and has since been recorded frc3m numerous localities 
in the Meditetrancan region, eastward to Greece, Turkey, 
Syria and Turkestan. Apparently most of the records 
outside the western Mediterranean range concern 
synanthropic localities, and the species is obviously 
easily distributed by commerce. Outside the palearctic 
region records arc still few, and since Jawlowski (1930) 
reported B. lusitatius from Mexico the species was only 
recently mentioned as occurring in the southeastern 
United Stales by Filka & Shelley (1980) where it seems 
to have been previously misidentilied as Brachyiulus 
pusillus (Leach). 

ft lusitanus is similar to B. pusillus, but differs in its 
slightly larger maximum size; it can be easily separated 
by the fanlikc phylacum of the posterior gonopods 
(spiniform in R pusillus) (see the illustrations in Verhocff 
(1910). Jawlowski (1930) and Strasscr (1976)). 




Records of millipedes introduced into Australia from 
elsewhere arc few. Baker (1978) extensively reported on 
the distribution and dispersal of the west Mediterranean 
julid Ommutoiuius tnotvleti (Lucas) in South Australia 
and Victoria. I have summarized the known records of 
European millipedes in Tasmania (Jeekel J981) and more 
recently added the oriental paradoxosomatid 
Orthoinorpha coarauta (De Saussure) to the Australian 
list (Jeekel l9S2a» 

The material of the South Australian Museum 
contained, besides some specimens of Ommatoiuius y 
also a number of samples of a Meditetrancan species 
of Brachyiulus not previously known to occur in 

Brachyiutus tusitanus Verhocff 

Brachyiulus pusillus lusitunus Verhocff, 1898: 153, PI 

6, Eig. 28 

Micrahrachyiulus calcivapus Verhocff, 1910; 225, PL 2, 

Figs. 33-34. 

Brachyiulus lushauus; Jawlowski, 1930: 183, big. 4; 

Strasser, 1976: 606, Fig. 37. 


Woodley's Vineyard (near Glen Osmond, SL 
Adelaide), soil, Berlese funnel, 30.VII-8.VI 1 1.1950, leu 
R. V. Southcott, 3c?, 59 , 8 juvs. 

Woodley's Vineyard, soil, Berlese funnel, 3. IX. 1950, 
leg. R. V. SoutheotU 4 juvs, 

Scllick's Milt, 23. IX. 1954 (E.S.I. 1953). leg. Ci. K 
Cross, I* , 

Ommutoiuius moreleti (Lucas) 

Zulus tuotvleti l.ucas, I860: %. 

Arcivutu.s uwreleu; Attems, 1928: 291, PI. 18, Figs. 


Si -hizophylluui mureleti; Sehubarl, 1966: 23, l-igs. 12-20. 

Omniatoiuitis moretetii; Baker, 1978: I. 


North End Pt. 1 incoln, I3.XU955, leg. M. Carrick, 

2 v. 


This species originates Irom the Iberian Peninsula, 
where it apparently occupies a rather restricted range 
in the nonhcrn half of Portugal and adjacent Spain. 
It has been reported from a number of Atlantic islands 
(Canary Islands, Madeira, the Azores, Bermuda, St. 
Helena), the Cameroons and South Africa. In Ausiralia 
it is known to become more and more wide-spread in 
South Australia and Victoria (Baker 1978) and it ha* 
also been reported from Tasmania, although as yet il 
does not seem to be so well established there. 

Onee it is introduced into a suitable environment it 
spreads rapidly and becomes a dominant clement of the 
millipede fauna of the area. On Ieneriffe (Canary 
Islands) for instance, it is seen almost anywhere in areas 
under human influence, Sehubarl (1966) gflVe an 
extended report on its distributional aspects in South 
Africa, where it appears to have spread over a large part 
of the country in the course of a few decades. Baker 
has described a similar phenomenon in South Australia. 

Ommatoiulus and Brachyiulus are easily distinguished 
from the autochthonous jnliform millipedes of South 
Ausiralia by the longitudinal striation of the dorsal side 



of the metasomites and by the freely projecting cauda, 
the latter being particularly distinct in Omtnato'tulus. 
Brachyiulus is further distinguished by having two 
yellowish dorsal paramedian bands. 


ATTEMS, C 1928. The Mvriopoda of South Africa. Arm. S. Ajr. 

Mus. 26: 1-431, pis. 1-26. 
ATTEMS, C. 1944. Neue Polydesmoidea. Zoo/. Anz 144: 223-251. 
BAKER, G. H. 1978. The distribution and dispersal of the introduced 

millipede, Qmmutoiultts moreletii (Diplopoda, lulidae) in 

Australia. I ZooL, Umd. 185: l-ll. 
CHAMBERI IN, R- V, 1920. The Myriopoda of the Australian region. 

Buff. Mus. comp. Zoo/, Harv. 64: 1-269. 
FILKA, M. E M and SHELLEY, R. M. 1980. The milliped fauna of 

the Kings Mountain region of North Carolina (Arthropoda: 

Diplopoda). Britnleyana 4: 1-42. 
JAWLOWSKI, H. 1930. On European Diplopoda introduced to 

America, tragm. fuuti. Mus. zoo/, po/on. 1: 183-185, pi. 5. 
JEEKEL, C. A. W. 1968. On the classification and geographical 

distribution of the family Paradoxosomatidae (Diplopoda, 

Polydesmida): i-vii, 1-162. Amsterdam {privately printed). 
JEEKEL. C, A. W. 1979. Notes on the classification of some liltle- 

known Australian paradoxosomatid genera (Diplopoda, 

Polydesmida) J. not. Hist. 13: 649-658. 

JEEKEL, C. A. W. 1980. A new genus and two new species of the 
family Paradoxosonuuidac from Australia (Diplopoda, 
Polvdesmida) Buli zoo/. Mus. Univ. Amsl. 7: 1-10, 

JEEKEL, C. A. W. 1981. Australia Expedition 1980; legit C. A. W 
Jeekel and A. M. Jeekel-Rijvers. List of collecting stations, 
together with general notes on the distribution of millipedes in 
eastern Australia and Tasmania. Vers!, techn. Cegev. Inst, faxo/i. 
Zoo/. Amst, 30: 1-59, 

JEEKEL, C. A. W. 1982a. Millipedes from Australia, I 
Antichiropodini from South Australia (Diplopoda, Polydesmida 
Paradoxosomatidae) Bu/f. zoo/. Mus. Univ. Amst, 8; 121-132. 

JEEKEL, C. A. W. 1982b. Eour new or little-known Australiosomatini 
from New South Wales (Diplopoda, Polydesmida 
Paradoxosomatidae) Bijdr. Dierk. 52; H 137-154. 

KARSCH, E 1881. Zum Studium der Mvriopoda Polydesmia. Arch 
Naturgesch. 47(1): 36-49, pi. 3. 

LUCAS, H. I860. Myriapodes, In "Notice sur I'histoire naturellc des 
Acores" (Morelet, A.), pp. 96-97. Bailliere, Paris. 

SCHUBART, O. 1966. Diplopoda III, Pselaphognatha 
Opisthospermophora, Colobognatha. S. Afr. anim. Life. 12: 9-227 

STRASSER, K. 1976. Uber Diplopoda-Clulognatha Grieehenland^ 
II. Rev. suisse Zoot 83: 579-645. 

VERHOEEF, K. W. 1898. Ueber Diplopoden aus Bosnien 
Herzogowina und Dalmaiien, IV. Theil: JuJidac. Arch. Naturgesck 
64(1): 119-160, pis. 5-6. 

VERHOEFF, K. W. 1910. Uber Diplopoden. 11-15. Aufsatz (31-35) 
Beit rage zur Kenntuis der Glomeriden, Jutiden 
Ascospermophoren und Lysiopetaliden, sowie zur Fauna Siziliens, 
etc. Nova Acta Acad Caesar: Leop. Carol 92: 139-448, pis. 1-9. 




by David C. Lee 


A study of sarcoptiform mites from surface soil (greatest depth usually = 4cm) at 9 florally diverse 
sites in South Australia is continued. Opisthosternal shields, supernumerary leg setae, gnathosternal 
fissures and cheliceral spatulae are considered separately. The Cryptostigmata is treated as 
including 5 suborders: Palaeosomatida, Retrofissurida, Afissurida, Profissurida and Comalida. The 
Afissurida and primitive Comalida (i.e. Mixosomatina and Clinofissurae) are considered further in 
order to complete the lower oribate mites within this study. 





South Australian Museum, North Terrace, Adelaide, South Australia 5000. 

(Manuscript accepted 20 December 1983) 


!_l I , D. K. 1 1 >kv Soiu.ptiUii-mcs (ALjri) of South Australian soils. 
4. Primitive oribate miles (C'ryplovtigmaia) wuh an eMrnsivc. 
unli\suivd hvstuanoial shield and aptychoid. ftec, S. Ausi. Mus. 
V){4): 39 6?". 

A study of sarcoptiform mites from surface soil 
(greatest depth usually 4 cm) at nine florally diverse sites 
in South Australia is continued. Opisthostemal shields, 
supernumerary leg setae, gnathosternal fissures and 
cheliceral spatulae are considered separately. The 
Cryptostigmata is treated as including five suborders: 
Palaeosomatida, Retroftssurida, Afissurida, Profissurida 
and Comalida. The Afissurida and primitive Comal ida 
(i.e. Mixosomatina and Clinofissurae) are eonsidered 
further in order to complete the lower oribate mites 
within this study. Five families are discussed although 
not represented by specimens in this study: the 
Eulohtnanniidae to confirm its inclusion in the Mixo- 
somatina; the Malaconothridae, Trhypochthonidae, 
Trhypochthonielhdae and a new family, Allonothridac, 
to delineate the two infraorders (Mixosomatina and 
Holosornatina) of the Comalida. Seven species of 
Afissurida, Mixosomatina and Clinofissurae were 
collected. Four species are new: Plarynothrus 
brevisetosm, Cromnia jethurmerae, Nanhermannia 
grandjeani, Phyllhermunma eusetosa. Two subspecies are 
new Epilohmannia cylindrica media, Epitohmannia 
pallida australica. Papitlacants pseudoaaculatas is newly 
recorded from Australia. A new synonym is Vepracarus 
under Papitlacants. New combinations are egypika and 
mdica (ex Paptllacarus) with hjhniannia, and koreanus, 
ogawai and ramirezoe (ex Vepracarus) with Papillacaras. 


This publication is a further part of an ongoing study 
(Lcc 1981, 1982). Its intent is to complete the systematica 
of what are generally regarded as the primitive, lower 
or macropyline oribate mites. Because the higher 
classification has changed so much since this study 
began, the oribaie mites considered here are not all 
closely allied. They appear to be similar because they 
are primitive but have an extensive unbroken 
nysleronotal shield (a derived character state convergent 
in two suborders) and are aptychoid (a character state 

ancestral to the derived ability to fold the proterosoma 
back onto the hysterosoma which has arisen in three 
suborders). The mites to be considered include the 
Lohrnannioidea (Afissurida), the Mixosornaiina 
(Comalida) and the Clinofissurae (Comalida - 
Holosornatina). The following families from the 
Mixosomatina have already been dealt with: 
Phthiracaridae and Euphthiracaridae under Euplyctima 
Lee 1981) as well as Gehypochthoniidae under 
Monofissurae (Lee 1982). 

The numerous higher categories of oribate mites may 
be reduced, some becoming superfamilies, if there is 
compensation for the disproportionate tendency to 
"upgrade" the classification (Krantz 1978). However, 
these categories are retained here. I follow the recent 
discarding by O'Connor (1984) of the abnormal use of 
"cohort" in acarology. Instead, between infraorder and 
superfamily, the rank of section is used as for 
crustaceans (Kaestner 1970). 

The mounting procedures used have sometimes 
resulted in squashed specimens. The possibility of such 
distortion, always represented in illustrations of 
gnathosterna where it facilitates description, should be 

All mites described in this study are deposited in the 
South Australian Museum, Adelaide. 


Opisthostemal Shields and Setae 

The opisthostemal shields may be defined by the setae 
they bear. Unfortunately, in the reduction from the 
primitive three pairs of setal files, it is difficult to assess 
which files are lost or merged. Therefore, the genital and 
anal shields are regarded as the movable shields covering 
the relevant orifices and bearing setal files Jg and Zg 
or Ja and Za; and if these shields bear only one setal 
file it is referred to as either JZg or JZa whether or not 
it appears likely that a specific file (e.g. Ja in some cases) 
is missing. The more lateral shields are the aggenitai 
shield bearing setal file Sg and the adanal shield bearing 
setal file Sa, which may be merged to form the 
optsthoventral shield. In the Retrofissurida and 
Afissurida, the anal shield may be split into a peranai 
shield (bearing one or two setae Ja) and a paranal shield 

■ I 

Kl I . S. At IS 1 MI'S !<) (4); V),(y' 

Mow r,<\- 

(bearing two, three or four setae la). In such cases, i 
have in the past (Lee 1982, Figs. 7 and 16) referred to 
these setal files with a different notation. In exceptional 
eases, as Hoplophthiracarus sheaisi Lee, 1981 (sec f ig. 
25), the movable shield over the genital orifice bear* 
three* seta] Hies indicating it is a merged genital and 
aggenual shield- 

Chaetotaxy of Legs and Coxites 

The leg tarsi of some species described below are 
hypenrichous. Lying proximally to the normal 
complement setae, the e.xlra supernumerary setae are 
restricted to the four following positions: two 
dorsolateral — da, dp and two ventral — «/v, pw The 
suffix c *jp* is added lo the signature for each 
supernumerary setae, followed by a number indicating 
the position of the including whorl or rank, with H \ n 
Lhe most proximal. 

I he \cntrnl cuxite setae are in approximately single 
file and are numbered as such from the adaxial end 
without any attempt to homologizc them with the files 
(anterior, median, posterior) from which Ihey might 
be derived. 

Gnathosternai Fissures and Cheliceral Spatula 

Gnathosternal fissures are regarded as derived and 
may be functionally correlated with greater rigidity of 
the idiosoma requiring greater mobility of the 
mouthparts. The two types of fissure have been defined 
(Lee 1984). The species described below have mentu- 
coxal fissures, whilst what was referred to as "diandric" 
(Orandjean 1957) or a "quadrangulale mentum ,v (Lee 
1982) occurs in more advanced taxa with dicoxal 
fissures, I hese two types of fissures split different parts 
of the gnathosternum (although the homologies of 
areas regarded as derived from either the palp eoxtle 
or mentum are noi confidenlly held), whilst before 1 he 
differences were regarded as due to a change in shape 
of the mentum. Apparently in rare cases both types 
of fissures occur on lhe same animal (see Nothrtn 
species. Tragatdh I93la: Hg 6), 

As previously (Lee 1984), the term cheliceral spatula 
is used for "TragSrdh's organ". The presence of a 
cheliceral spatula was regarded as a possible 
synapomorphy for the Holosomatina (Corualida) (Lee 
1984). Norton (personal communication 1982) has 
since pointed out that the cheliceral spatula is present 
in a number o\' families of Mixosomatina (Comalida) 
and I discuss this further below under ''Remarks*' on 
that infraordcr. 


I accept the concept that the Ctvptostigmata is a 
paraphyletie taxon as suggested by O'Connor (1984), 
with the Astigmata regarded as a sister-group lo part 
of the Comalida (in the restricted sense used below). 

On the other hand, I do not in tins instance follow the 
principle of Wiley (1981) (hat paraphyletie groups be 
rejected as unnatural, arbitrary, human constructs. I 
consider it uselul to retain the Cryptostigmata as a 
grade, sharing similar primitive character states, and 
"left behind" by the considerable differentiation of the 

Thc higher classification of the Ctyptostigmata IS 
modified so thai it is regarded as including five 
suborders, sequenced to reflect their supposed level of 
advancement as follows: Palaeosomatida (sec 
Bifemorata Lee, 1981), Rctrofissurida (see Ketrofissurae 
Lee 1982), Afissurida (see Afissurina Lee, 1984 and 
heluw), Profissurida (see Profissurae Lee, 1982 and 
Pediculochehdae) and Comalida (see Comalida, in 
part, Lee, 1984 and below). Changes from the 
classification presented at the 6th International 
Congress of Acarology in 1982 (Lee 1984) are that the 
Dismalida is disbanded and the included three distinct 
primitive lineages upranked to suborders, whilst the 
Profissurina is excluded from the Comalida and also 
upranked to a suborder. Four of these suborders may 
be regarded as elades, but the Comalida is not a 
complete clade since the Astigmata is excluded from 
it (although it may prove arguable that the entire 
Comalida is a sister-group to the Astigmata). 

Upranking of the Profissurida is a response to lhe 
proposition (Norton, O'Connor and Johnston 1983) 
that the Pediculochehdae and Haplochthoniidae 
constitute the latest derivative lineage within the 
Ptotoplophoroidea ( = Profissurida). Even if this 
phylogeny was well supported, I would mainlain the 
Pediculochehdae as a separate superfanuly on the basis 
of morphological divergence (widely separated coxites 
lll-IV from III, terminal anus, pretarsa] stalked pad), 
although 1 do tentatively accept that with the 
Protoplophoroidea it constitutes a monophyletic group. 
On the other hand the late derivative phylogeny of 
Pedieulochelidac is not acceptable since there is no 
evidence suggesting an ancestor with hysteronotal inter- 
calary sclerites and erectile setae, or that the absence 
o( sclerotuation and reduced sctation (often a 
convergent similarity) are synapomorphies for a lineage 
composed of it and the Haplochthoniidae. I propose- 
that the Pediculochehdae is regarded as an early 
derivative group as indicated by the primitively disjunct 
externa! malae. This suggests that the conjunct external 
maiae oi other Profissurida have arisen separately as 
an apomoiphy convergent lo a similar character state 
in the uiHJoi ity of the Comalida and therefore supports 
the upranking and exclusion of the Profissurida from 
the Iumct suborder 


D/^//o^/'v.-Livptosuginata. Hysteronotal shield lacks 
transverse FftSurcs but transverse furrows often present. 
Hysteronotal gland absent. Hysteronotal ehaetotaxy 



holotrichous or hypertriehous. Seta] lilex wilh two seLac, 
both similar in size and shape to other proteronoial 
setae. Coxites merged, forming two podostemal shields 
(coxites 1, 1) or Ml, IV), Chelieeral spatula abseni. 
ExteroaJ malae disjunct. Gnathostcrnal fissures absent, 
but mentocoxal furrow sometimes present Adoral setae 
in transverse row; at least adaxial two pans large and 
flattened. Prelarsi with one claw. Tarsus I and genu 1 
each wi|h (wo solenidia. Tibia 1 solenidia flagelliform. 
Femora undivided. Nymphs with similar fades to adult. 

Remarks: The Afissurida, also referred to as 
Afissurina (Lee 1984), includes only one superfamily, 
the Lohmannioidea; Grandjean, 1969, I he 
Lohmannioidea was then included in the Mixonomata 
which approximates to the Mixosomatina (Comalida) 
considered below. Despite this, Grandjean (1950), in a 
thorough work on the Lohmannidae, has indicated thai 
this family is very distinct from other oribatc rnites, but 
with distant affinities to certain "Enarthronota". I agree 
that it shares a number of primitive character states with 
the Retrofissurida (part of old *Enarthronota") such as 
disjunct external malae and a complete natal chaetotaxy, 
including two large, well-separated setae in file s and 
sixteen pairs of hysteronotal setae. On the basis of this, 
it is reasonable to regard the absence of a hysteronotal 
gland as primitive and not secondary as in some 
Mixosomatina (Comalida). Therefore, my inclusion of 
the Lohmannioidea in a separate, primitive suborder. 
with its cxicnsivc hysteronotal shield considered as 
convergent with that state amongst the Comalida, 
reflects the earlier (1950) conclusions o\' Grandjean 
rather than his later (Grandjean 1969) work. 

Besides the Lohmanniidae, two other families have 
been recognised in the superfamily, each including only 
a .single species. The Nothrolohmannidae Balogh, 1968 
is recognisable by bifurcate seta A humeral processes 
bearing seta 51 and large triangular aggenital shield 
fused to posterior margin of coxite IV. The 
Xenolohmauniidae Balogh and Mahunka, 1969 has a 
* l meiuon divided into two parts, not meeting medially'*. 
As illustrated, this structure appears as if it might 
represent a pair of broad Hat processes extending 
forward from coxites 1 rather than an unusual 

Family LOHMANNIIDAE Berlese 

Lohmannini Berlese, 1917b: 176 
I ohmanmidae- Grandjean, 1950; 100 
Lohmanniidae: Wallwork, 1962a: 457 
Lohmanniidae: Corpuz-Raros, 1979: 31 S 

Type-genus: Lohmannia Michael, 1898: 75 

Diagnosis: Afissurida. Preanal shield present, either 
bar-shaped, T-shaped, or reduced to central knob. 
Aggenital shield sometimes present, but not fused to 
coxite IV or extending posteriorad to halfway along 

genital shield. No broad flat process extending forward 
from coxiic t under gnathosternum. Hysteronotal seta 
51 noi on conspieuous humeral process, Proieronotal 
SCta /I not bifurcate. 

Morphology: Small to large (480-900) dull yellow or 
brown mites. Proteronotum long (0.4x length of 
hweronotum) and broad (as wide as hysteronotum). 
Opislhosternum usually .shorter lhan podostcrnum. Legs 
short (leg ] longest, less than 0.4x idiosornal length). 
Palp femur and genu fused or partially fused. Genital 
shield bears 10 or more setae in two files (usually ftsfe 
4Zg) % may be divided into subequal anterior and 
posterior pans. Setal files Sg and Sa absent. Anal shield 
may he divided into peianal and paranal pans. 
Solenidiotaxy I (2-1-2), II (1-1-1 or 2), 111 (1-1-0), IV 

Distribution: Widespread in tropics, where greatest 
generic diversity occurs. Particular species may 
themselves be widespread, Achieved "a greater degree 
of evolutionary radiation in Africa than in South 
America" with "a wide distribution in those parts of 
the world which belonged, formerly, to Gondwanaland'* 
(Hammer and Wallwork 1979). fn temperate regions, 
but generic diversity decreases away from tropics, 
Lolvnannia being most widespread, tn northern 
hemisphere, genera other than Lohtnunnia only found 
south o!*45 e N. 

Some species known to burrow in roots or bark. 
Represented both in deserts and moist habitats such as 
coastal bogs. Ecological notes on species collected in 
Philippines (Corpuz-Raros 1979) indicate wide variety 
of niches which family can occupy in limited 
geographical area. 

Remarks: The above diagnosis distinguishes the 
Lohmanniidae from the two very similar monospecific 
lamilies also in the Afissurida and is, therefore, relatively 
trivial. A number of further character states are listed 
under "Morphology" which may apply to all three 

Balogh (1972) included 20 genera in the 
Lohmanniidae. Amongst these, authors have regarded 
the following four genera as similar to Papillacarus 
(which is represented in this study): Cryptacarus 
Grandjean, 1950; Dendracarus Balogh, I960; 
Lepidacarus Csiszar, 1961; i'epracarus Aoki, 1965b. On 
the basis of the disposition of the opisthosternal shields 
Cryptacarus and Dendracarus can be excluded. Whilst, 
as elaborated on below, Leptdacants can also be 
distinguished, Vepracarus is regarded as synonymous 
with Papillacarus. 


PupillacarvsKunsi, 1959: 70. Type designation (original): 
' ' lohmannia mun hides Berlese v. aciculata Beilese 


Rl ( . s \nsi. \irs_ im ,i»_ v>->,,- 

Ma\\ WH5 

i'epmcartis Aokj, 1965b: 142. Type designation 
(original): " Veprucarus ogauvi Aoki v specuov!— o.syn. 

Type-species: Papillacarus aciculatus (, 1905: 24). 

Diagnosis: Lohmanniidac. Genital shield divided by 
transverse fissure into Two subequal shields. Anal shield 
divided by longitudinal fissure into peranal and paranal 
shields, Preanal shield reduced to central, bieomate 
knob. Rostral tectum blunt T anterior margin either 
smooth are or sinuous with four slight tubercles or with 
three conspicuous tubercles (lateral tubercles where 
lamellae meet margin)- Proteronotal plasraic seta (z2) 
with long cilia on only one side of slim or lanceulatc 
main stem. Notal setae with cilia which may be 
inconspicuous or so long that setae bush-like, Posterior 
notal hypertrichy, never anterior to seta 72. Notal 
integument papillate. Ventral ridges on femora I and II. 
Analehaeiotaxy: 2Ja> 47m, Tarsi III and IV each with 
10 setae. Tarsus II witJi two solenidia. 

Distribution. Possibly almost limited to Old World. 
Virgin Islands (NTa); Annobon Island, Ghana (Hw); 
Crimea, Moscow, Uktainia (Pe); Bulgaria, Greece, lialy> 
Spain, Tunisia (Pm); Korea, Japan (Pe); India (Oi); 
Thailand (Os)l Java* Philippines (Orn): South Australia 
(Aa). Known Oriental fauna shows greatest 
morphological diversity. 

Found in grassland, feeding on grass roots, also under 
trees, where grass may or may not have been present. 

Remarks: Three genera (l^epidacarus, Papillacarus, 
and Veprucarus) have the disposition of opisthosternal 
shields given in the above diagnoses, tepidacarus can 
be distinguished from the other two genera by the 
preanal shield being an undivided central knob, as well 
as by the hysteronotal setae being leaf-like. 

The only species of Papillacarus and Vepntcarus that 
has been described in enough detail to include characters 
of the gnathosoma and legs is R angutatus. Therefore, 
characters used to delineate genera and their 
included species have to be limited to those of the 
idiosorna and in some cases to the notum. On the basis 
of such characters 1 have not been confident about 
distinguishing the two genera. Instead I have merged 
them and regarded Papillacarus as including the four 
following species-complexes based on the form and 
chaetotaxy of the hysterouoial setae. This is intended 
as a temporary measure until more characters are 
considered. Although the aciculatus-complcK (includes 
type of Papillacarus ) and f/^nw/-eomple\ (includes type 
of Veprucarus) are easily delineated, this is undermined 
by the other two complexes. The fact that the hirsuius- 
complex includes species from both genera illustrates 
this confusion. The type complex is considered last since 
it includes the species found in South Australia. P. 
egypricus Elbadry and Nasr, 1977 and P. indicus Hafl fc2 
Kardar, 1972 are grouped in Lohmunnia, nx'iimb. 


Diagnosis: Papillacarus. Hypertrichy on hysteronolum 
posterior to seta ./4 or 75, area bearing 50-60 sciac 
Supernumerary setae short (length subequal to or less 
than distance between posterior setal bases), bush-like 
(because long cilia)- Complement setae similar, 
indistinguishable, if any longer seta (uncertain if super- 
numerary or complement), then around posterior 

Remarks: Three species included in the complex, R 
hirsutus (Aoki, 1961), ex Cryptacar'us — Pc r Om, Ap. R 
chamartinesis Perez- 1 nigo, 1 967— Pm. R koreanus 
(Mahunka, 1973), ex Veprucarus, rueonib.— Pc. 


Diagnosis: Papillacarus. Hypertrichy on hysteronorum 
posterior to seta /4. area bearing 50-80 setae. Super- 
numerary setae short, bush-like. Complement setae at 
least twice as long, with inconspicuous cilia. Single rank 
of tubercles between setal ranks 5 and 6. 

Remarks: Two species included in complex P ogawai 
(Aoki, 1965b), ex type of Veprucarus, nxomb. — Os. R 
ratvirezae (Corpus- Raros, 1 979), e\ Veprucarus, 
nxomb.— Om. 


Diagnosis,' Pupdlucurus. Hypertrichy on hysteronolum 
posterior to seta 72, area bearing 120-170 setae. Super- 
numerary setae short, bush-like, Complement setae J4 t 
and those posterior to it, more than five times as long, 
with inconspicuous cilia. 

Remarks: Two species included in complex. P navosus 
Balogh, 1961— Om. f?m/^t'Corpu/-Raros, 1979— Om. 

aeiculatus-com plex 

Diagnosis: Papillacarus, Hypertrichy on hysteronotum 
posterior to seta J4, area bearing 20-45 setae. Super- 
numerary setae short but similar in form to complement 
setae, so both with inconspicuous cilia, not bush-like 

Remarks: Six species included in complex. P. 
aaatlatus Berlesc, 1 90S — Pe. Pm. P angulatus 
W'allwork, 1962a— Bw P nndirositvlus Aoki, l965b-Os. 
R ondriast Mabunka, 1974 — Pm. P. pseudoaciculaius 
Mahtinka, 1980a— Pm, Aa. P v/tis Elbadry and Nasi, 
1977— Pm. 

Papillacarus pseudoaciculatus Mahtinka 
(Figs 1, 2) 

Papillacarus pseudiiaciculatus Mahunka, 1980a: 126 

Dull, ochre-coloured with yellowish brown eheliceral 
extremities, external malae and legs. Cuticle of shields 
mainly granulate except in transverse hysteronotal 
furrows and in snntll, usually marginal patches. Similar 

v\K< niMimUMlMM AKM) Ol sol.' Ill \i STRAI » AN SOU S 4 

but less extensive areas papillate, protuberances bCJUg 
aeieulate (as few on gnathosternum, illustrated Fig, 1), 
Cuticle of prehystcronotal fissure strongly striated 
suggesting propodosoma can flex downward. Idiosomal 
length 510 (1); appendage lengths— ch 42.5, pa 45, / 180, 
// 160, Iff 135, fV 175; femur breadths— //</ 12.5, / 50, 
//45, W37.5, H 42 5. 

Chelieeral seta ch\ much shorter than chl. 
Vlentoeoxal furrow present, clearly not fissure. Rostral 
icel urn hyaline* indistinct, possibly smooth arc. 
Dorsolateral longitudinal fissure from anterior margin 
of hysteronotal shield back (o level of preanal shield. 
This delineates dorsal margin of pleural shield which 
bears no setae but slit-like pore (hf\) at anterodorsal 
corner. Few super numeral y hysteronotal setae (?six") so 
area behind ,/4 bearing about IS setae. Cilia on 
hysteronotal setae longer than depicted on type 
specimen, more like P. ondnasi (Mahunka 1974: 575, 
Fig. 8), but supernumerary setae relatively longer. Seta 
/6 conspicously longer than surrounding setae. Six 
transverse hysteronotal furrows, only anterior furrow 
jusi belli ud seta J\ complete, posterior furrow level with 
seiH 74 reduced io two indistinct pits. Indistinct 
structures may represent two pairs ol genital papillae. 
Rive pairs of setae on anterior genilal shield and four 
pairs tin posterior genilal shield (Ug. 2Z#), possibly 2g3 
missing from adult complement. 

Material examined; One tritonymph (N 1983 1), bases 
of grasses and plantains, Glcnthorne, 12.6.1974, D. C. 

Distribution: Tunisia (Pm); South Australia (Aa). 
South Australia: Glenthorne, cuinvaied pasture, I tnto- 
nymph ( /?>. 

Remarks: I he referral ol tins single specimen to be 
a tritonymph o\' R pseudoacicuiams is not done with 
confidence. The identification is based mainly on the 
low number of supernumerary hysteronotal setae, but 
this may be related to it being an immature stage. The 
extensive soinal coverage with aeieulate protuberances 
in dense patches excludes some other species. Since the 
species are so similar in the acicutatus-complzx, the 
above tritonymph could represent a new species. 

Suborder COM A LI DA 

Diagnosis: Cryplosligmata. Hysteronotal shield 
usually lacks transverse fissures (exceptions: TBI present 
ill most Lumixosornatae. } .T8) present in some 
Platynothmssud Croumia species). Hysteronotal gland 
usually present (exceptions, absent in Eulorunanniidae, 
Nanhermanniidae, Phthir-aearidae, Synichotrithnae). 
Hysteronotal chaetotaxy usually hypotrichous, wiih at 
least seta J4 and/or /A vestigial or absent (holotrichous 
exceptions: some b'umixosomatae, Eulohmannhdae, 
some Clinolissurae). Setal file s with one or two setae, 
if two, then at least one 0.5x length or less, of 
protcronotal $etei /2. Coxites rarely discrete, varying 
degrees or lusion may merge into single podosternal 

<hie!d. Chelieeral spatula present or absent. I xternal 
malae conjunct or coaretate. Griathostemal fissures 
absent or menlocoxal and/or dicoxal fissures present . 
Adoral setae positioned so that at least ao\ anterior to 
ao2. Pretarsus wirh one, ihree or rarely two claws. Genu 
1 with one (true for all species with two solenidia on 
tarsus I), two or three solenidia I ibia I solenidia 
baculiform, piliform or flagelliform. femora undivided. 
Nymphs with similar or dissimilar iacies to adull. 

Remarks: The Comalidu (Lee 1984) is diminished to 
exclude the Profissurina as commented on under 
"Sysicmalics" above, This leaves it restricted to iwo 
infraorders: Mixosomatina and Holosomaliua. Fhe 
Mixosomatina and amongst the Holosomaiina, the 
Clinofissurac, are dealt with below. Thus, within mv 
study of South Australian oribate mites, lIucc sections 
(Phcrenotae, Gymnonolae and Potonotae) of 
Holosomatina will not have been considered. 

The synapomorphy o\" the Comalida is still xl 
as the conjunct or coarctaic position of the external 
malae, but it is now treated as convergent with that 
character state in the upgraded Profissurina A fenetat 
trend can be recognised in the adults to derived character 
slates such as a strongly integument, fused 
somal shields, fissured gnathosternum, conspicuous 
soma! ridges (lamellae, pteromorphs, teeta), fewer hairs 
(setae, solenidia), heteromorphic leg segments and a 
more complex respiratory system (poies, uaehcaej. Since 
a similar trend does not occur in the immature suu;e\, 
they lend to he dissimilar to the adults in L il! but the 
more primitive taxa. 

Relationships wilhin I he Comalida, which includes 
the great majority of extant oribate species, arc not well 
understood. I have assigned the advanced taxa to the 
Holosomatina, which is regarded as monophyletic I he 
remaining distantly related taxa arc provisionally referred 
to the Mixosomatina. which is regarded as a primitive 

The Comalida is regarded us paraphylctic because the 
Astigmata is not included (sec "Systcmatics" above). 
This is preferred since it provokes the search amongst 
the Comalida for the sisier-grotm 10 the Astigmala, a 
more acceptable phylogeny, since the primitive nature 
ol ParliypodulitmiiissuvgcM^ that it is ancestral within 
a lineage including both o\' these ta.xa. If this search is 
not fruitful, however, it may prove desirable to regard 
the entire Comalida as a sister -group to the Astigmam 
and so monophyletic. 


Diagnosis: Comalida. Transverse midpodosKiual 
fissure between coxites II and ML Often two solenidia 
nn gfcriU L if only one solenidium then cither transverse 
hysteronotal fissure TBI present i-.-.m, 
Gehypochthoniiclac) or opislhostemal setation reduced 
(at least setae Sg absent and usually only one seta JZu— 


Kl < S ,MIS1 Ml IS )*) (4) W-o? 

Iter. W 

Neomixosomatae). Adana! .shields nevei merge behind 
anal shields- Chelieeral spatula present or absent. 

Remarks: The Mixosomatina includes three sections 
( Eu m i xosom a tae, Mcsomi \osom a lac and 

Neomixosomatae) much as before (Lee 1984) except that 
the latter section has been extended to include (he 
Trhypochthoniidac and Trhypochthoniellidae. Within 
my study of South Australian oribatc mites, members 
of the Eumixosomaiac have already been considered 
under "Monofissurae" within the "Arrhronotina'* (Lee 
1982), but it should be noted that the recently included 
(Lee I9S4) North American Nchypoehlhoniidae Norton 
and MctA 1980 lacks a transverse hysteronotal fissure, 
which requires a change in the diagnosis. The other two 
sections arc considered below, although no members of 
the Neomixosomatae were found in this sludy. 

The Mixosomatina lias no syuapomorphy, but is 
recognisable by the primitive character state of a 
podoMernal fissure which allows rhe propodosoma to 
move in relation to the inctapodosoma. In some taxa 
with a strongly sclerosed cuticle, litis flexibility is 
accentuated by a number o\' derived slates such as in 
piychoidy, when the legs are pulled up into ihc soma 
and the proteronotum flexed downward to close on a 
forward facing section of the genital shield, or when the 
propodosoma can partially telescope into the meta- 

Relationships within the Mixosomatina are uncer tain. 
partly because some primitive character states may or 
may not be secondarily reverted to in advanced taxa. 
The weakly selerotized eulicle of the Eumixosomatae 
is primitive in the Parhypochibioidea and the 
Nehypochthoniidae but in the Neomixosomatae it may 
be either reverted to or there may not have been any 
strongly sclerosed ancestors. Furthermore, whilst t (Lee 
1984) considered that the absence of chelieeral spatulae 
was primitive but diagnostic of the Mixosomatina, 
Norton (personal communication, 1982) has pointed out 
that chelieeral spatulae do occur in some Mixosomatina. 
He has observed chelieeral spatulae in Coltohmarwia 
gigantea and a new North American species of 
Collohmanniidae. Also, their presence in 
Perlohmanrhidae is suggested in an illustration by 
Grandjean (1958: Fig. 5A), and I have now included the 
Trhypochthoniidac and frhypochthonielliclae in 
Mixosomatina, both of which have chelieeral spatulae. 
Tragcirdh (1931b) recorded chelieeral spatulae on 
Phtfuracarus maculatus, but this was refuted by 
Grandjean (1959) who considered the structure to bean 
oneophysis. Chelieeral spatulae are considered to be 
absent in all Eumixosomatae, some Mesomixosomaiae 
(Epilohmanniidac, Eulohmanniidae, Phthiraearidac, 
Lupthiracaridae) and some Neomixosornaiac 
(Malaconoihridae). Therefore, although primitively 
absent, some taxa without chelieeral spatulae may have 
ancestors that possessed rhem. 

The three sections within the Mixosomalina arc 
g fades. The Mesomixosomaiae include a number of 
specialized lineages, whilst the Eumixosomatae are 
apparently primitive (ceitainly so in the case of 
Parhypaclithontits, but possibly Elliptochihonius reflects 
specialization to living in the deepei soil layers) and the 
Neomixosomatae include advanced lineages which may 
be ancestral to the Holosonialina, Regarding t he- 
relationships of the Astigmala, 1 hold (o a conservative 
belief thai weak scleroiizatiou in ihe Asrigmata is an 
ancestral character slate and not a reflection of ncoteny 
as proposed by O'Connoi (1984). Therefore, a sister- 
group to the Asiigniata would have a weakly scleroLized 
adult, as well as lacking a rostral tectum and chelieeral 
spatulae. Possibly, reduction in the fourth hysteronotal 
setal rank would be the synapomorphy, placing this 
sisler-group within the Eumixosornalae. However, a 
stringent analysis will be a necessary preiude to 
developing a durable model for the areas only speculated 
on here. 


Diagnosis: Mixosomatina. Hysteronotal shield 
without transverse fissures. Rostral tectum present. 
GnathosLemum v\ irh A -shaped mentocosal fissure 
Adoral setae in lha*e conspicuous pairs. OpislhostcmaJ 
setal files $g with at least one seia, J/a wiih a! least 
two setae. Protcronoial setal file $ with one or two setae. 
Genu I with two or rarely three solemdia. 

Remarks: The Mesomixosomatae is a diverse group 
of small families which* although having a strongly 
selerotized adult cuticle, have maintamed flexibility by 
developing various specialized ways of moving the 
propodosoma in relation to the hysterosoma. I have not 
attempted any superfamily groupings, Even grouping the 
six families into aptychoid (Collohmanniidae 
Grandjean, 1969; Lpilohmanniidae Oudemans, 1923; 
Eulohmanniidae Grandjean, 1931; Ferlohmanniidae 
Grandjean, 1954a) and ptychoid (Huphthiracaridae 
JacoL 1930; Phthiraearidac Petty, 1841) taxa is 
questionable. The ptychoid families have already been 
considered in this study (Lee 1981) as the Euptyctima, 
The Epilohmanniidac is represented in collections for 
this study and is considered below. Although not 
represented, the Eulohmanniidae is also considered in 
order to confirm its grouping in this section despite 
possessing some primitive character stales suggesting U 
might even be grouped with the Afissurida. 


Lesseriidac Oudemans, 1917: 78 
Lpilohmanniidae Oudemans, 1923: 79 
Lpilohmanniidae: Grandjean. |954a: 430. 
Epilohmanniidac: Grandjean, 1969; 144 
Lpilohmanniidae- Norton, Met/ & Sliarma. Wflf: U.\ 
lype-genus: Epilolununnia Berlese, 1917b, 

SAKC OJMlrokMt S (V.'AKI) 01 s« >l. Ml -M SIliM \\\ SOU S I 


Diagnosis: Mcsomixosomalac. Minute lo medium- 
sj^ed (320-800) yellow to brown miles. Intensive non- 
■scleroti/ed cuticle al transverse midpodostcrnal fissure 
between coxites II and III allows propodosoma to 
partially telescope in and oul of mctapodosama. 
Otherwise, extensive sclerotized somal shields, with 
coxites fused to each other, aggeuital shield and usually 
mentum (exception Epilohmannoides Jacoti). Posterior 
end dJ aggeuital and auteriot end of adanal shields 
truncated or fused together so that ventrolateral 
longitudinal fissure straight. Adanal shield merges 
medially as broad band in front of anal shields. 
Cheiieeral spatula absent. Palpal segments fused so thai 
only two separate. Adoral seta ao\ bifurcate. 
Hysleronnial gland present, poie opens into depression 
also containing alveoJus of relic seta ZA, whilst sela M 
similarly absent, Opisthosternal set a I file Ag includes 3-7 
pairs. Opisthosternal pores /<?/" and 5c// present. On 
tarsus I seta pd? reduced to scale closely associated with 
distal face of solcnidium so4. On tibia 1 solenidium 
flagclliform, longer than segment. 

Distribution: Possibly cosmopolitan. WallwOrk 
(l%2b) stales that Epilohmanniidae Appears to be 
widely distributed through the warmer regions of the 
world". Records from southern Canada (Nn) and 
Moscow (Pe) could be most northerly, with records from 
South Australia (Aa) being most southerly, suggesting 
predominantly pantropical distribution becoming sparse 
in temperate regions up to 57 °N and 35 °S. Outside Ant- 
arctica and Subantarclie, South Ethiopian and New 
Zealand-Australian may be only minor regions in which 
family not represented. 

Mierofmbitats range widely from grass roois, 
vineyards and forests, but possibly do not include arid 
or semi-arid environments. 

Remarks' The Epilohmanniidae includes the 
typegenus and Epilohmanmndes. Norton, Mei/ and 
Sharma (1978) in describing two species of 
Epilohmannoides also give d diagnosis lor the family 
on which the above diagnosis is based. 

1 PILOtlMANNlA Beriese 

Lptiohmamua Beriese, 19|7b: 176 lypc designation 

(original): "l.ohmanniu eylindrica BcrK 4 . 
Lesseriu Oudemans, 1917: 78. lypc designation 

original): by monmypy ( n Le$$etiil szanis/oi Oudms. 

1915", synonym of Lpilnhmannia cyfindriea by van 

dor Hammen 1959: 54), 
Ivpe species liptiohmanma cy/indnca tBerlese, 1905: 

Mavjiosis: Epilohmanniidae. Opisthovenlrai shield 
divided into two parts by transverse fissure just posterior 
to genital shield- tieniial and aual shields oblong rathei 
than suboval; posterior margin of genital shield and 
anietioi margin of anal shield being broad-shouldered. 
Trochanter III and IV with distal axis at right angles 

to proximal axis. Acetabulum IV on same longitudinal 
line as III, both divided by spurs into double opening- 

Distribution: As for Ppilobmanniidae. 

Remarks: The more complex form o\' trochanter 111 
and IV and their acebatula with double openings. 
suggest that these legs can lock into either of two hasie 
positions and, further, indicates that Epilohmattnia may- 
be derived from the much less diverse (four compared 
wiili 26 species) Epilohmannoides with simpler posterior 
trochanters and acetqbula. 

Balogh and Mahunka (1979) have distinguished 
within Lpilohmannia a subgenus, Sinolohmannia, bj the 
presence of a spine-like seta d associated with tlte 
solenidinm on tibia IV. The n7///r///<r/-eomplex of 
Schuster (I960) can be regarded as equivalent to the 
nominate subgenus, A more conspicuous character state, 
the position of the acetabulum on eoxite IV, has not 
been referred to in grouping species, Both the species 
considered below have the acetabulum IV on the 
posterior margin of coxiie IV, but it may be near the 
anterior margin or in between. However, the included 
species arc similar and, unfortunately, the specimens 
collected in this study belong to species with a number 
of even more similar subspecies. Since I am not 
confident about grouping them in any of the subspecies, 
new subspecies have had (o be established. Because oi' 
the similarity o\' the established subspecies, references 
to them are included under (he new subspecies. 

The illustrations of one species in this paper are 
attended to show character states delineating the family 
and genus rather than the subspecies. 

Epihhmaania cyfindriea (Beriese) media n.ssp. 
(figs 3 5) 

Lolinmunnia cvfindritv Beriese, 1905: 23. 

Lessetta s^misloi Oudemans, 1917; 78. 

Eptlohmannia szanishi Sehuslcr v I960: 202 (including 

minima, p. 205). 

Epilohmaania cyhndrica Aoki, 1965c: 309. 

Epilahmannia cylindriea Bayoumi and Mahunka, 1976; 



Dull (slight gieasy shine) chestnut brown, slightlv 
darker cheiieeral extremities and external malae, 
extensive pale si raw -coloured cuticle between 
protcrosorna and hvsterosoma when extended . Most of 
hystcrostcrnum wUh sparse puneta, limited in other 
parts of venter to areas illustrated (Fig. 3). Idiosomal 
length (abutting margins to midpodosomal fissure, 
possible extension 50, retraction 30) 465 (10, 415-525); 
appendage lengths (for 475) - eh 45, pa 50, / 195, 7/ 175, 
/// 195. /I ' 275; femur breadths—/*/ 7 5, / 20, // 20, /// 
ISi /I 20; broadest segment breadth— A genu 32.5, // 
genu 30, /// trochanter 25, IV trochanter 37.5. 

Appendage setae*, eh (2K pa 13 8), / (1-4-5-5-17), // 
(1-4-5-5 12), HI <2-3-4-4 10), I\ (2 3-4-4-9). Solcmdia; 
/ (2- Ml. // (1-1-2). IU (l-l ()> ( /I <!-!-()). Solcnidi.. of 


REC. S. AUST. MUS. 19 (4); 39-67 

May. t§85 

100 *j 


FIGS. 1-5. PapUlacarus and Epilohmannia; 1-2, PapUlacarus pseudoacicutatus Mahunka, tritonymph; 1, gnathosternum; 2, right chelicera, 
anterior surface; 3-5, Epilohmannia cylindrica (Berlese) media n.ssp., female; 3, idiosternum; 4, gnathosternum; 5, right chelicera, 
anterior surface. 

^XktOlM IIOKMI S (AC ,\Kl> oi- SOUTH m.-stkai \,\\ SOU S i 


genua and tibiae flagellitbrm with minute, closely 
associated seta d (possibly partially coupled— sec 
Epilohmannoides tetnie: Norton* Metz and Sharma, 
1978), on legs III and IV seta d only recognised when 
separated from solendium by preparation of specimen. 
Soleniclium so4 {so3 regarded as absent) on tarsus t also 
flagelliform with minute scta/^/3 closely associated with 
distal face. Other solenidia on tarsi I and 11 ccratiform. 

Chelicera without spatula, small process (Fig. 5) 
posterior to movable digit probably chilinous plate 
representing reduced trochanter. Adoral setae ao\ and 
au2 bifurcate. Genital shield setae: 5/g i3*g. 

Tarsus IV with length 5.5x breadth, and setae v] and 
v2 (see Schuster I960; Fig. 5 a- Id and A'2) spine-like, 
but less robust than far /?. cyiindrica minima (see 
Schuster I960: Fig. 5b). Seta \2 nearer to vl than to i'3. 

No eggs observed, specimens assumed female since, 
amongst eight pairs of setae on positor, three pg pairs 
well apart from rest, two dg pairs longer, One to three 
boli in each specimen, light to dark brown 
hetcrogcncously granular contents without recognisable 
cellular structures. 

Material examined: South Australia (Aa). Hololypc 
female (N 19832), nine paratypc females 
(N19833-NI983U), bases of grasses and plantains, 
Glemhortie. 12.6.1974. D. C. Lee. 

Distribution; E. cyiindrica (subspecies media only 
known from South Australia). Tennessee (Na); Tchad 
(He); Austria, Egypt, Hungary, Italy, Moscow, Sicily, 
Spain (Pe); Turkestan (Ps); Philippines tOm); South 
Australia (Aa); Hawaii (Ap). South Australia: 
Olenlhorne, cultivated pasture, 10 (2/8). 

Remarks: i rv/indrica media lies between E. c. 
cyiindrica and E c. minima in a gradation of character 
states, As indicated by Schuster (1960), IE c. cyiindrica 
is 435-573 long, has tarsus IV length 5.6s. breadth, with 
sciae i'I and vl setose, whilst E. c. minima is 370-440 
long, has tarsus IV length 3.6x breadth with setae vl 
and v2 robust and spine-like (more so than £ a media). 

Epihhmamua pallida Wallwork austmlka n.ssp. 
(Fig. none) 

Epilohmannta pallida Wallwork, 1962b; 689. 

Epikdimannia pallida pacifwa Aokl 1965t" 312. 

Epitohmannta pallida aegypdea tiayoumi & Mahunka, 

1976; 8, 

Epilohmannta pallida indica Bhattaeharya & Banerjee, 

1980: 19. 

Epilohmannta pallida americana Balogh & Mahunka, 

1981: 59, 


Dull brown, paler but similar to E. cyiindrica. 
Idiosomal length (abutting margins to midpodosomal 
fissure, possible extension 42.5. retraction 25) 400 (2 c\ 
Piccaiiiiime Ponds, 420-425, 3 c\ Chambers C Hilly, 

350-405); appendage lengths (for 405)— ch 40, pa 45, 
/ 153, // 130, /// 140, IV 190; femur breadths— /w 10, 
/ 22.5, // 20, III IS, IV 15; broadest segment breadths—/ 
genu 27,5, //genu 25, ///trochanter 17.5, /J'troehantei 

Appendage setae: ch 2, pa (3-8), / (1-3-4-5-15), // 
(1-4-4-4-12), /// (2 or 3-3-3-3 or 4-10), IV (2-3-3-4 9> 
Compared with E. c media seta \>a absent on femur L 
genua I arid III, whilst seta vp absent on genua III and 
IV. On genu I seta r small. Solenidia similar in number, 
shape and si?e to £ c media, except on genua III, IV 
and tibiae HI, IV, solenidia relatively longer. 

Chehcera without spatula. Adoral setae aa] and aol 
bifurcate- Genital shield setae: Xlg, }Zg, 

Characters previously used to delineate subspecies as 
follows, Hysteronotal setae sparsely ciliate and tapering, 
sclaZI posterior to 71, distance VIZI subequal to 71-Jl. 
Anterior apodeme to coxite 1 curves back to single 
apodemc separating coxites I and II without connecting 
ridge to partner, coxile seta /// 2 subequal in length to 
/// 3, lateral margin of coxite JV .straight. Opisthostcrnul 
seta Sg2 level with Jgl, setal file Zg separate from file 
Jg by distance subequal to length of Zg\, slit pore Saf 
inclined at angle less than 25 D from transverse axis. 
Distal end of trochanter III without dorsal spur, on 
tarsus IV seta vl nearer vl than v3. 

No eggs observed, specimens assumed female since 
amongst eight pairs of setae on positor. Three pg pairs 
well apart from rest, two dg pairs longer. One or two 
boli in each specimen, light to dark brown 
heterogeneously granular contents without recognisable 
cellular structures. 

Material examined; South Australia (Aa). HoloLype 
female (N1983L2) and two paratype females (N1983I3, 
N1983I4), grass and moss or litter under Eucalyptus 
viminali\ Chambers Gully, 12.6.1974, D. C lee. Two 
paratype females (NI98315, N1983I6), litter and sparse 
grass under Acacia sophorae, Piecaninuie Ponds, 
20.8.1975, D. C. Lee. 

Distribution; E. pallida (subspecies ausrralica only 
known from South Australia). Texas, Minnesota, North 
Dakota, South Dakota (Na); Paraguay (Nib); Ghana 
(L'w); ij'gypt tPe); West Bengal (Oi); Hong Kong (Os); 
South Australia (Aa); Hawaii (Ap), South Australia: 
Chambers Gully, savannah woodland, 3 (2/8); 
Pieeaninnie Ponds, coastal closed-scrubland 2 (2/8). 

Remarks; E. pallida is not very different from E. 
cviindrica (especially if the comparison is made to fi 
c minima) except that seta dl on tarsus IV (5 
conspicuously spine-like. 

The other five subspecies of £ paliidanrc very similar. 
/: /? austialica can be distinguished from these by the 
character states listed above. On the basis of these il is 
most similar in /;. p. indica and E /?. pacifica t bur dWfcr^ 
in two states regarded as important; the shape of the 
anterior coxite apodemes and the position of seta \'2 
on tarsus IV. One character, the chactotavy of the genual 


Kits AIM. MMS I') Hi M& 

\/av, Mffi5 

shield sometimes used ro distinguish autepttici. has not 
been listed as such because of its intraspecific variation 
(see MeDaniel and Bolen 1983). 

Family EULOHMANNIIDAE Grandjean 

Eulohmanniidac Grandjcan, 1931: 144 
Eulohmanniidac: Grandjean, 1954a: 429 

lype-genus: Eulohmamua Berlese, 191 

Diagnosis: Mesomixosomatac Medium -si?ed 
(650-700) straw-coloured mites. Extensive nonsderotized 
cuticle at transverse midpodostcrnal fissure between 
coxites II and III allows propodosoma to partially 
telescope in and out of metapodosoma. Otherwise, 
extensive scleral ized somal shields, with coxites fused 
lo each orhcr and aggcnital shield. Posterior end of 
aggenital and anterior end of adanal shields taper so 
that ventrolateral longitudinal fissure V-shaped. Adanal 
shield not merged medially in front of anal shields. 
Chelieeral spatula absent. Palpal segments fused so four 
separate. Adoral seta ao\ not bifurcate. HystcroTiotal 
gland absent. Hysteronotal chaetotaxy holotrichous, 
fourth seta! rank not reduced, Opisrhostcrnal setal file 
&g hyperlrichous and dispersed, 15 or more setae. 
OpisthoMcrnal pore &//' present, ^/absent. On tarsus 
[. seta/?tf3 setose and well separated from solenidium 
so4. On tibia I, solenidium piliform, shorter than 

Distribution; Holaretic. 

Remarks: The Eulohmanniidac includes one species, 
Euiohmonmu ribagai Berlese, which has been described 
a number of times, perhaps the most accurate and 
comprehensive illustration being that by Lebrun and 
Wauthy (1981: Fig. 2). But there appears to be no 
detailed description o^ the gnathnstcrnum. Since the 
species has a number of primitive character states such 
as a holotrichous hysteronotal chaetotaxy and no 
hysteronotal gland, it was necessary to check the 
gnathostemum to evaluate whether or not the species 
was correctly grouped in the Comalida. 1 his was made 
possible by a gift of two specimens from Dr. Georges 

Eulohmmnia ribagai Berlese 
(Fig. none) 

I.ohmanma (Eulohmonmu) ribugui Berlese, 1910: 223. 


Idiosomal length 655 and 695. Chelieeral spatula 
absent. Mentocoxal fissure present, A-shaped. External 
malae conjunct, distally with two lateral robust refract ile 
teeth and median hyaline flap. Gnathosteraal chaetotaxy 
reduced: 3fcq 2n>, ]pa Gnathostemum elongate much 

as Perlohtnunniu dh.sunilis (see Woolley 1969: Fig. 10). 
Tarsus \ with 23 setae and lliree solenidia. The shape 
and positioning of plasmic seta d\ and three solenidia 
much as Perlohmunnia dissimilis (see Grandjean 1958: 
Fig. 6C); seta d\ short with distal knob, solenidia well 
separated from serae. 

Material examined Two adults (N1983I03, N1983104), 
litter and humus, Lauzella Wood, Belgium, 4J9S0* ft 

Remarks: Having seen details of the gnathostemum, 
there is no doubt that Eulohmannia is well placed in the 
Comalida, and Ihercfore that some apparently primitive 
character states, such as the absence of hysteronotal 
glands and holotrichous hysteronotal chaetotaxy, are 
derived reversals. Although Eulohmannia in general 
torm and extent of sclerotization resembles 
Epilohmannia, it exhibits similarities to Perlohmannia 
of the gnathostemum and tarsus I which suggests they 
may be regarded as sister-groups. 


Diagnosis: Mixosomatina. Hysteronotal shield 
without transverse fissures. Rostral tectum present. 
Gnathostemum either without fissure or A-shaped or 
what may be transverse linear mentocoxal fissure 
present. Adoral setae present or absent. Opisthostcrnal 
scum 1 file Sg absent, JZa usually with only one seta 
(exception: Mueronoihnts—UZa). Proteronotal setal file 
$ absent, plasmic seta z2 may be reduced and seliform. 
On genu I, one solendium. 

Remarks: The Neomixosomatae was established (Ire, 
1984) to include the Malaconothridae which has a 
conspicuous fissure between coxites 1 1 - 1 1 1 and lacks a 
chelieeral spatula, but is in some ways similar to the 
Nothroidea, the most primitive of the Holosomaiina. 
As indicated in the above "Remarks" under 
Mixosomatina, it lias now been established thai 
chelieeral spatulae occur on some of its members. This 
means that the Trhypoehthoniidac: Balogh, 1972 is not 
excluded from the Mixosomatina by possessing 
chelieeral spatulae and so the relevant literature has been 
examined, even though no representatives were collected 
in this study. The result is that a number of changes 
are made to the classification, most genera of 
Trhypoehthoniidac including the type-genus being 
included in the Mixosomatina. On the other hand, two 
genera have been grouped in a new family retained 
within the Holosomatina. The three families 
(Malaconothridae, Trhypoehthoniidac and 
Trhypochthonietlidae) now in the Neomixosomatae arc 
briefly commented on below. 

1 wish to emphasize the provisional nature of this 
classification and that I have ignored some character 
states considered in the more comprehensive studies of 
relevant families by Knulle (1957) and van der Ham men 
(listed by van der Hammen, 1959) because they are only 

SAKI IMMIIOKMl S I At Akll iW 5l H I I 1 1 At IS I KAl IAN MUI S ■» 


known lor a lew species. Since those studies, lour genera 
have been added to the eight genera (included by 
Balogh, 1972 within the Trhypochthonhdac and 
Malaconothridae) and, both before and after the studies, 
ihe majority ul' species descriptions have not been 
extensive enough, iu some cases with debatable generic 
combinations. Therefore, a thorough reinvestigation of 
established ta.\a is needed before reliable diagnoses can 
be given lo them, and Lneir considerable relevance to 
the classification of ComaLida understood. 

Family MALACOMN HRIDAK Berlcse, 1917b 

Dm^nosis: Neomixosomatae Cheliccral spatula 
absent, Adoral setae reduced to two pairs or all absent. 
Onalhostcrnat fissure absent or, if part uf mcntoeoxal 
fissure present, lateral parts never meet at mid-point. 
Proteronotum without setal file s and seta rl setiform 
and less than 0,5x length y2. Hysteronotal setaZ2 nearer 
Z3 than J2. Loxite II posterolateral corner extended as 
carina with backward facing socket. Opisthosternal setal 
file Sa with three setae. On tarsus I, three solenidia 
clustered between level of setae (XR-pdi. On tibia I, one 

Remarks: "I lie posterolateral corner of coxite II may 
act as a socket into which trochanter III fits, possibly 
functioning as a pivot when the propodosoma flexes to 
one .side. Such a movement could be limired in some 
species by a large tectum, projecting laterally from jusl 
posterodorsally to acetabulum II, hitting a similar 
smaller tectum anterior to acetabulum III. 

The Malaconothridae, as by Balogh (1972), includes 
the following four genera: fossonodirus Hammer, 1962; 
Mulaeonothms Series^ 1905; Trimataeonudirus Bcrlese, 
191 7c; Zeanothms Hammer, 1966. 

Family TRHVPOCHTHONIHMK Willmann, 1931 

Diagnosis; Nenmixosomatac. Cheliceral spatula 
present. Adoral seiae with three pairs present 
Ylculocoxai fissure present, A-shapcd. Proteronotum 
with seta z2 club-like oi, if set i form, either subequal in 
length 10 Jl or, il less than 0.5x length. A seta s present. 
Hysteronotal seta 21 nearer 72 than Zl. Coxite II 
posterolateral corner sometimes extended as carina with 
backward facing socket. Opisthosternal setal file Sa with 
(wo or three setae. On tarsus I, rhrec solcnidia usually 
widely spaced between level of setae pc/2-pd4 (exception* 
ftydronoihrtis). On tibia f usually two solcnidia 
(exceptions: Hydn)tiotlirn\ and Mtterotiothrus with one 

Remarks: The eight genera included by Balogh (1972) 
in the Irhypocluhoniidae are decreased in number by 
subdivision either into the Irhypochthoniellidae or the 
Allonothridae (Clinofissunie). I he following foui 
genera are si ill included .-t/Wwo^Vtis Grandjean, 1931; 
llydronothnts \oV i , 1 964; Mucmnothrus fragandh. 

19? 1c; Trhypoehdwnias Berlese, 1905 Ihe conservative 
emphasis on notal setae results in llydronoihrus being 
grouped \n (his family, although the leg [ setation is 
similar to that of the Malaconothridae. Archegozetes 
differs considerably from the other three genera, 
especially with regard to its long leg tarsi and seta t2 
But some recorded differences, the absence of botli a 
cheliccral spatula and a fissure between coxites II and 
III (Beck. 1967), are regarded as errors. The presence 
of a cheliccral spatula and a coxite 1 1/1 11 fissure was 
recorded by Gtpndjean (1959) and van der Hammcu 
(1955) and has been confirmed by Dr. R. A. Norton 
(personal communication. 1983) on specimens from 
Mexico, Panama (NTm), Brazil (NTb) and Malaysia 


Diagnosis: Neomi.xosomatae. Cheliceral spatula 
present Adoral setae with three pairs present. 
Gnathosiernal line (?fissure) present, transverse, linear. 
Proteronotum vvjih seta :2 club- like or set) form and less 
than 0.5 x length j2, Hysteronotal seta Zl nearet ./2 than 
73. Coxite II posterolateral corner not extended as 
carina. Opisthosternal setal file Sa with two setae. On 
tarsus 1, three solcnidia widely spaced between level of 
setae pd2-pd4. On tibia I, one or two solcnidia. 

Remarks: Trhypoehthontellidae includes two genera: 
f'rhypoehthoniellus Willmann, 1928; Afronodvus 
Wall work, 1961. The grouping together of these genera 
is mainly based on an assumption about gnulhosternal 
siruclure They can be regarded as having a quadrate 
menium; a M hypostome reehteckig" for 
Irhypoehthonietlus (Knullc (957: 151) or as illustrated 
for AJronothrus (Wallwork 1961: Fig. 7, Hammer 1972: 
Fig. 19a). The line illustrated on Afronodirus could not 
be the dico.xal Rssure which delineates Ihe anterior 
margin o\ the "quadrate mentunV 1 oi advanced 
Holosomatina. It is therefore assumed that it is a 
mentocoxal fissure and there may be a valid quadrate 
mentum, but careful evaluation ofwhai the drawn line 
represents is still needed. 


Dta^nosis: Comalida. Coxites fused together into a 
single shield. On genu I, one solenidium. Tibia 1 
solcnidia baculifonn, piliform or, if fiagcllifotm, closely 
issocialcd with similar seta. No transverse hysteronotal 
fissures (exception; 7V/JI present in some Crofonia and 
Plarynothrus species). Usually two setae in file JZa t 
always when adanal shields not merged behind anal 
shields. Cheliccral spalula present. 

Remarks: The Holosomatina is regarded as 
nionophyletie, whilst its four subordinate sections 
(Clinofissurae, Phereuotac, Ciymnonotac and 
Poronotue) are likely to be grades. Ihe lineages have not 


ki •: s. \i s i mi is im in }« ft 

My. /W* 

been identified, although the Poronotae (being the most 
derived) could be a clade. Only the most primitive 
section, the Clinofissurac, is considered in detail below 
\ major la.xon, Lhe Circumdehisccnliae Grand jean, 
|9S4a ( = either the Euoribatida Balogh and MaluiTika, 
1979 or a major pan of the lkachypylina: Batogh, 1972 
or the Pherenotae. Gymnonotae plus Poronotae) >s 
regarded as monophyletic but remains unnamed in this 
classification as before (Lee 1984). This is because if 
approximates to the Holosomatina (excluding only the 
six families of the Clinofissurac) and because the fusion 
of coxites into one shield (which is the synapomorphy 
of the Holosomatina) is regarded as a predominant step 
that precludes conditions such as plychoidy and is a 
prelude to oilier stares correlated with a thick, rigid 
idiosomal integument, such as a circular hysterosomal 
dehiscence line, a dicoxai gnathosternai fissure and a 
Tracheal system opening ventrolateral^' between legs II- 
1 1 1 or in the acetabular cavities o\' legs I or 111. The 
presence of a dicoxai gnathosternai fissure may be 
correlated with the circular hysterosomal dehiscence line 
of the "Circumdehiscentiae", but a Nothrus species is 
clearly described by Tragardb (1931a) as having both 
a dicoxai and a mentocoxal gnathosternai fissure, 
although this needs confirmation since it is difficult to 
conceive the function of such a double-jointed system. 
The presence of a tracheal system appears lo be the 
synapnrmorphy of a slightly smaller taxon than the 
' ( Cireumdchiseentiae ,t since it is absent from the 

Section CMNOriSSDRAi: 

Diagnosis: Holosomatina. Gnathosternum usually 
wiih A -shaped mentocoxal fissure present (exceptions: 
no fissure in Allonothridae, possibly both mentocoxal 
and dicoxai fissure present in one Nothrus species). If 
adanal shields separate from hysteronotal shield (i,e, 
exclude Nanhcrmamiiidae) they do not fuse behind anal 
shields. Genua subquadrangulate in outline and similar 
in size lo tibiae. Hysterosomal dehiscence line T-shaped, 

Remarks; The Clinofissurac is diagnosed by character 
states primitive to the Holosomatina. It is equivalent 
to the majority of the Nothroidea plus the 
Nauhermannioidea and Hermannioidea in the previous 
classification of Balogh (1972), when the latter two 
superfamilies were included in the higher oribate mites 
(as *"Brachyp\iina l, K although they are excluded from 
the similar Circumdehiscentiae Grandjean, 1954a. On 
the other hand, this is the first lime that members of 
the Nothroidea have been included amongst the higher 
oribate mites (if considered as referring to the 

The following six families are included in this Section: 
Allonoifrridae n.f.; Carnisiidae Oudcmans, 191)0; 
CrotoniidaeThorell, 1876; Hermanniidae Sellnick* 1928; 
Narihermannitdae Sellnick,. 1928; Nothndac Berlese, 

1885. All but the Allonothridae and Nothridac are 
represented in this study and the Allonothridae is 
considered further as it has to be defined. The 
Allonothridae, Carnisiidae, Crotoniidae and Nothridae 
can be grouped in the Nothroidea GTandjean, 1954a* 
but superfamilies will not be considered here. 


lypc-genus; AHonothrw van dcr Hammcn, 1953 

Diagnosis; Clinofissurac-. Gnathosternai fissures 
absent, or, if part of mentocoxal fissure present, lateral 
pam never meet at mid-point. Three pairs of 
adoral setae. Rostral tectum without median incision. 
Proteronotai plasmic setae jZ at least 2x as long as 
distance jl-<X and sctiform or slightly swollen distally. 
Hysteronotal seta M absent. Hysteronotal gland present. 
Notal setae z\ and ./5 not on apophyses. Coxite selae 
not hypcrtrichous (3-1-3-3). Setal file Sg abscnl, 
combined setal file JZg on median margin of genital 
shield, fdiosorna not almost covered in continuous 
shield. No separate preanal shield. Palp tarsus with nine 
setae. No dorsolateral supernumerary setae on tarsus I, 
Nymphs without small shields around hysteronotal setal 

Remarks: The Allonothridae includes the following 
two genera: Allonothrus van dcr Hammcn, 1953; 
Pseudonothms Balogh, 1958. 1 he genera are similar to 
each other and were included in the Trhypochthonhdae 
(Neomixosomatae, see above), but because o\^ the 
apparent fusion of all the coxites into one shield they 
are now grouped in the Holosomatina as a new family. 
The Allonothridae exhibits similarities io both the 
Neomixosomatae and Clinofissurac suggesting that they 
belong to the same lineage, but much more data is 
needed before a ciadistic classification can be proposed 
within the Comalida. 

Family C4MISIIDAK Oudcmans 

Carnisiidae (part) Oudemans, 1900: 142. 

Carnisiidae (part): Sellnick, 1928: IS. 

Carnisiidae: Grandjean, 1954a: 431. 

Carnisiidae (part); Sellnick and Forsslund, 1955: -J73. 

Carnisiidae: van dcr Hammcn, 1959: 65. 

Type-genus: Camisia von Heyden, 1826 

Diagnosis: Clinofissurac. Gnathosternai A -shaped 
mentocoxal fissure present. Three pairs of adoral setae. 
Rostral tectum without median incision. Proteronoial 
plasmic seta zl may be reduced and globular but never 
enclosed in bothridium, if filamentous length 0.75x-1.5x 
distance j2-z2, Hysteronotal seia 74 present or absent. 
Hysteronotal gland present. Notal setae ;;l and J5 
sometimes on apophyses. Coxitc setae not hypertrichous 
(3-1-3-3 or 4). Coxitc shields usually not merged with 

SVi( nnilUkMLN i.V.AKHOI SOL !M U SI KAI I \N SO 1 1 N 4 

: I 

aggenital shields. Setal file Sg includes two selae T 
combined setal file JZg on mediau margin of genital 
shield. Idiosoma not almost covered in continuous 
shield Discrete preaual shield usually as wide as anal 
shield- Palp taisus with seven setae. Tarsus I with four 
or fewer dorsolateral supernumerary setae Nymphs 
without small shields around hysteronotal setal bases. 

Distribution: Possibly cosmopolitan, greatest diversity 
in temperate regions, represented in tropica] montane 
or oceanic regions. 

Remarks, Camisiidae are small to gigantic (530-1225) 
dull brown mites, usually covered in cerotegument, thick 
in parts and with adhering detritus and fungus. 
Camisiidae has been considered as synonymous with 
Nolbridac, but both names have been in use since 
Grandjean (1954a) separated rhem. The family is now 
still as regarded by Balogh (1972) except that 
Austronoihms is grouped in Crotoniidae (Ramsay and 
Luxton 1967), Comprehensive descriptions have been 
made for Camisia (Behan 1978; Andre 1980) and 
Plotyno flints (Fujikawa 1982). 

The following four genera are included in Camisiidae: 
Camisia von Heyden, IK26; Hemtnoihrus Berlesc, 1914; 
Neonothrus Forsslund (in Schick and Forsslund 1955); 
Platynothrus Berlesc 1914. Heminothrus and 
Neonothrus are very similar to Platynothrus, 
Heminothrus being established earlier (p. 38) in the same 


Platynothrus Berlesc, 1914; 99 Type designation 
(original); "Nothrus palliatus K. <-/V bistriatus K )'* 

Type-species- Platynothrus peltifer (Koch, 1839: 29/9), 

Diagnosis: Camisiidae. Protcronotal plastnic seta ^2 
vermiculate, usually slightly dilated and ciliate distally. 
One seta in file s, Seta si without conspicuous apophysis 
(at most, shorter than 0.5x distance cl-^l). Bothridial 
cavity with only one side pocket situated near base of 
seta z2- Hysteronotal seta J4 absent or minute. Seta J5 
without conspicuous apophysis (at most, shorter than 
its own diameter). Two setae in file Sg on inner margin 
of aggenital shield. T\vo setae in file JZa Coxites 1, 11, 
ill merge across midsternal line, coxites IV may be 
partially or completely separated from each other by 
fissure. Leg I with six solcnidia (1-2-3). 

Distribution: Widespread within temperate regions, 
usually montane or oceanic in tropics. Canada, 
Greenland (Nn); Argentina, Bolivia, Chile, Peru (NTc); 
St. Helena (t:s); Finland, Norway, Sweden, U.S.S.R., 
other parts of Europe fPeJj Japan (Ps): Himalayas (Oi); 
South Australia (Aa); New Zealand (An); South Georgia 

Remarks: Major works on Platynothrus are included 
in studies on the Swedish fauna (Sellick and Forsslund 
1955: 513), Bcrlese^s collection (van der Hammcn 1959: 
71), Himalayan collections with a key to species (Aoki 
1965a; 290) and the northern Japanese fauna (Fujikawa, 
1982: 279), Balogh (1972) regards Plutynothms as having 
one pretarsal claw so his keys do not work for species, 
such as the one described below, which have three 
pretarsal claws. Heminothrus and Neonothrus are very- 
similar to Platynothrus, and might be grouped in this 
genus. Platynothrus includes seventeen species and two 
of these have a subspecies. 

PUttymtthrus brevisetosus n.sp. 
(Figs 6-12) 


General appearance and measurements: Red-brown, 
covered in cerotegument, rhiek with adhering dctrtlus 
and fungus laterally and posteriorly on hystcrosomaand 
proximally on legs. Notal minute pits and lateral low 
bumps distributed as illustrated (Fig. 6), whilst all shields 
covered with fine punctuation except on rostral tecrum. 
Legs similar sculpturing, mainly proximal, ventral and 
sometimes anterior on individual segments. Setae, claws, 
external malae, chcliceral digits clear or light brown and 
refractile. Idiosomal lenglh 1140 (1); appendage 
lengths— ch 65, pa 100, / 580, // 490, /// 470, /K615; 
femur breadths— pa 25, / 75. 11 75, /// 65, IV S2.5. 

Prosternum: Lateral refractile half of external malae 
in vertical plane when unsquashed (\jc. not flattened as 
illustrated, Fig. S) and bears two rows of cilia on dorsal 
surface. Adoral setae aoi blunt-ended. Setae pel and c2 
in shallow, punctated depressions. Coxites all fused into 
one shield, but clearly delineated by grooves, and 
separate from aggenital shields. 

Proteronotum: Seta z2 vermiculate, but slightly dilated 
and ciliate distally. Bothridial cavity with only one side 
pocket just median to base of seta z2 y so posterior 
median wall forms unbroken arc. Surface has central 
flat-topped mound with two mounds behind it, and 
mounds above acetabula for legs. 

Opisihosfernum: Opisthoventral shield separate from 
coxites, without protrusions from inner margin to bear 
Sg setae. Prcanal shield abuts more closely on to anal 
shield than illustrated (Fig. 7) and appears foreshortened 
since extends upward in vertical plane. Opisthosternal 
selae in file Sa blade-like, with hyaline flaps on setal core. 
Genital shield with anteriorly twisted downward median 
margin bearing selae (Fig. 7) rather than median ridge, 
also transverse central unpigmented /one, Twenty-three 
marginal JZg setae flattened with slightly ragged, blunt 
ends. Pores Z<a/and Saf present. 


HEC -S. AUSI, MUS. 19 (4); 39-67 

Mtt\\ I9H5 



B 9 

FIGS. 6-9. Phtynothms brcvheiost/h n.sp.. female; 6, notum; 7, uliosiemum; K. gnathosternum; *), right cheliecra, anterior surface. 



Hysteronotum: Fissure runs transversely along first 
setal rank and backward along setal files Z and S (SI, 
52, Z3, ZA) possibly representing longitudinal pleural 
fissure Sclae 72, Z2, 73 lie between median ridge and 
lateral furrow. Many bysieronotal settle appear bladelikc 
with hyaline Haps to setal core and marginal cilia, not 
always illustrated (Fig. 6). Seta 74 apparently present 
but rainule. 

Appendages: Chelicerac relatively small. On fixed digit 
five teeth including distill point; anterior and posterior 
teeth in two pairs, Seta t7;l originally present, but both 
losl during dis-section. Setae: ch (2), pa (1-0-3-7), / 
(0-10-5-6-28 or 29), // (1-10-5-6-23), Ul (4 or 5-6-5-6-23), 
/K(0-5-4-6J3). Soleniclia: pa (0-0-1), / (1-2 3), /f (1-1-2), 
/// (I 1-0), I) T (1-1 -0). Pretarsus with three subequal claws, 
lateral claws with inconspicuous dorsal cilia file 
Terminal plasmic setae on palp tarsus spine-like, in 
recess. Solenidia baculiform, relatively short never 2x 
length of associated seta, longest on genu 1. Tarsi all 
with supernumerary setae. Setae colourless, on tarsi have 
strongly refractile bases but hyaline tips. 

Somal inclusions: No eggs or boh, Ovipositor 
involuted, if extended would be about 3s length of 
genital shield. Breadth slightly less than that of one 
genital shield. Bears eight pairs of setae, two ~dg pairs 
longer than setae JZg> three pg pairs more than length 
of genital shield away from three nig pairs. 



Material examined: Holotype female (N198328), litter 
under Finns pinea, Knott Hill Forest, 22.5.1974, D. C. 

Distribution: Souih Australia— Aa: Knott Hill, 
cultivated pine forest, 1 (1/2), 

Remarks: R brevisefosus, with three pretarsal claws, 
short hysteronotal sclae and well-spaced setae ./2, keys 
to P. albmontanus Hammer, 1958 from Bolivia in Aoki's 
(1965a) work as would R tenuicluva Hammer, 1966 from 
New Zealand which is possibly even more similar. & 
btevisetosus is distinguishable from these two species by 
the presence of a rudimentary 74, hysteronolal fissures, 
large size, blade-like hysterosomal setae and stout setae 
j\ and z\. 

Family CROTONIIDAE Thorcll 
Crotonoides Thorcll, 1876: 452-508. 
Holonothridae Wallwork, 1963; 727. 
Crotoniidae: Ramsay and l.uxton, 1967: 479 

Type-genus: Crotonia Thorell, 1876. 

Diagnosis: Clinofissurae. Gnathosternal A -shaped 
mentocoxal fissure present. Three pairs of adoral setae. 

Rostral tectum without median incision. Proteronotal 
plasmic seta c2 reduced, globular and enclosed within 
bolhiidium. Hysteronoial seta ./4 present. 
I lysteronotal gland absent. Nolal sclae t) and 75 usually 
on conspicuous apophyses Casite / or // setae 
sometimes hypenriehous. but not III and IV (3 or 4 - 
I or 2-3-2 or 3). Coxite shields merged with aggerulal 
shields. Setal file Sh includes two or three serae, 
combined seial file JZg on median margin o\' genital 
shield, Idiosoma not almost covered in continuous 
shield. Discrete preanal shield as wide as anal shield. 
Palp tarsus with nine selae. Tarsus / with five pairs of 
dorsolateral supernumerary setae. Nymphs with small 
shields around hysteronotal setal bases. 

Distribution: Southern temperate regions as for 
Crotonia. Austronofhrus from New Zealand, 
Holonothnts from New Zealand and Maequaric Island 
(An, Sa>. 

Found in plant litter, and on lichen, liverworts, moss, 
ferns and above-ground parts of shrubby plants. 

Remarks: Crotoniidae are large to gigantic (801-1500), 
dull, dark brown mites, extensively covered in 
cerotegument and detritus which may be accompanied 
by nymphal skins and fungi. The unique structure of 
The hoihridium enclosing a globular z2 is regarded as 
the family synapomorphy. 

The following three genera are included in 
Crotoniidae: Austronothrus Hammer, 1966; 
Ho/nnorhtm Wallwork, 1963; Crotonia Thorcll, 1876. 


Uesiwoodia Piekard-Cambridge. 1875. 383-390. lype 

designation (anginal): "Wcstwoodia obtecta sp.novT 
Crotonia Thorcll, 1876; 452-508. Type designation 

(original): "Crotonia obtecta (Pickard-Cambridge, 

Acronothrus Herlcsc, 1917a: 65. Type designation 

( or iginal): ki Nothrus (Acronothrus) cophinarius 

Mich. [908?*; 

type-species: Crotonia obtecta (O. Pickard-Cambridge, 
1875: 386). 

Diagnosis: Crotoniidae. Hysteronotal setation reduced 
to thirteen pairs (72, 73, Z\ missing). Bases of setae 71 
and Z.2 without connecting ridge. Rostral tectum with 
single prominence bearing both setae j\. Setae z,\ on 
separate conspicuous apophyses. Setae 75 (and other 
posterior setae) on separate apophyses or on single 
median branched apophysis. Posterior margin of coxites 
IV in smooth arc not broken by median notch. 
Opisthosternal file JZa includes three setae. 

D/sfribution: Widespread in southern temperate 
regions (NTc; Ee, Es; Aa, Ap, An; Sa)— see Hammer 
and Wallwork (1979) for a review, Note that South 


REC S. AUST. MUS. 19 (4): 39-67 

Mav. IP85 



11 \ pd 






pv2 ^ 


FIGS. 10-12. Pluiy/wthrus breyisetosus n.sp., female; 10, leg I, dorsal surface of genu, tibia and tarsus; 11, leg tarsi, posterior surface of distal 
ends; 12, leg III, posterior surface of trochanter and femur. 



Ethiopian record is St. Helena. More recent records from 
Tierra del Fuego (Mahunka 1980b) and New Zealand 
(Luxton 1982). 

Usually found in moist forest or heath in the plant 
litter, moss or on the above-ground parts of shrubby 
plants (Luxton 1982). 

Remarks: The thick layer of cerotegument, detritus 
and nymphal skins obscures the dorsal features of 
Crotonia mites as illustrated by photograph of C. obtecta 
(O. Pickard-Cambridge) (Ramsay and Luxton 1967: Fig. 
8). Crotonia may currently include twenty-four species, 
and Luxton (1982) provides a key for fifteen adequately 
described species. 

Crotonia jethurmerae n.sp. 
(Figs 13-19) 


General appearance: Dark brown, thick cerotegument 
with adhering detritus (including charcoal, left by forest 
fires), fungal hyphae and tritonymphal integument. 
Encrustation forms anterior protrusion encompassing 
setae z\ and posterior protrusion encompassing setae 
74, 75, Z5, S5 (small on some specimens, then bilobed). 
Notal minute pits and low bumps distributed as 
illustrated (Fig. 13). Setae, claws, external malae, 
cheliceral digits clear or light brown and refractile. 
Idiosomal length 1400 (3, 1390-1405). Proportions of 
appendages similar to those for male holotype (see 

Prosternum: Lateral refractile half of external malae 
in vertical plane when unsquashed (i.e. not flattened as 
illustrated, Fig. 15) and bears two rows of cilia on dorsal 
surface. Three adoral setae, ao\ bifurcate, flattened, one 
branch forming denticulate hyaline flap. Coxites all 
fused into one shield, partly delineated by grooves but 
leaving broad, flat mid-sternal zone, fused to aggenital 
shields. Hyaline cap on coxite seta may be broken off, 
leaving slim setal core. Coxite setae IIB and IV\ on 

Proteronotum: Seta z2 globular, enclosed in bothridial 
cavity with faint reticulate markings on lining, and 
appears as if slit-like opening to exterior between pair 
of ear-like folds. Cuticular pits confined to concavities 
at sides of high, flat median zone, which falls steeply 
to rostral prominence. No setae observed in file s, but 
if small would be difficult to distinguish amongst 
adhering detritus. Apophysis to seta z\ medium length 
(about equal to distance z\-z\) with curved lateral Hap. 

Opisthosternwn: Aggenital shield fused to coxites and 
only narrowly joined to adanal shield, with raised 
median rim near preanal shield and small notch anterior 
to seta Sgl (one female has 3Sg on one side). Preanal 
shield well separated from other shields, foreshortened 
as illustrated (Fig. 14) since extends upward in vertical 

plane. Number of opisthosternal setae blade-like, with 
hyaline flap on setal core. Genital shield has central 
transverse unpigmented zone embracing setae JZgS, 
JZg6 and notch between them (structure suggests shield 
may fold along this line). No pore Zaf located on anal 
shield although Saf and fa/3 conspicuous. 

Hysteronotum: Fissure runs transversely along beside 
first setal rank and backward along setal files Z and S 
(SI, 52, Z3, Z4) possibly representing dorsolateral 
longitudinal fissure. Ventrolateral longitudinal fissure 
terminates anteriorly, dorsal to posterior margin of 
acetabulum IV, so that anterior part of opisthosomal 
pleural shield merges with podosomal shield. 
Comparison with Camisiidae, adult and nymphs, 
suggests seta "dV is Z2 not 72, so hysteronotal 
ehaetotaxy 4./, 5Z, 4S, but concept of centralward 
migration of Z2 and lost 72 debatable. 

Appendages: Chelicerae relatively small, both digits 
terminating in paired, nearly parallel teeth (including 
distal points), with single large tooth proximally. Setae: 
ch (2), pa (1-1-3-9), / (0-12-5-6-38), 7/(0-12-5-6-30 to 32), 
77/(5-7-5-5-28 or 29), IV (1-8-5-5-29 to 31). Solenidia: 
pa (0-0-1), / (1-1-2), // (1-1-2), 77/ (1-1-0), IV (1-1-0). 
Pretarsus with three subequal claws, lateral claws with 
dorsal cilia file. Four terminal plasmic setae on palp with 
minute distal knob. Solenidia piliform, never more than 
1.25x length of associated setae. Solenidia on genua and 
tibiae II, ///, IV similar to one on genu I (Fig. 17), but 
similar-sized or smaller than associated setae. Tarsi all 
with supernumerary setae. Setae pale brown, or colour- 
less on tarsi with strongly refractile base but hyaline 

Somal inclusions: Three paratype females each with 
four or five eggs. Eggs smooth, suboval, 305-310 long, 
170-175 broad. Ovipositors involuted, extended would 
be about 2x length of genital shield. Breadth slightly 
less than that of one genital shield. Difficult to make 
out setae, setae dg longer than 7Zg. Two clearly 
delineated boli present per specimen, mainly cellular 
material, spores, hyphae, cellular sheets of plant tissue. 


Measurements and spermapositor (otherwise as 
female): Idiosomal length 1185 (8, 1125-1290); 
appendage lengths (for 1185, holotype) — ch 77.5, pa 110, 
/ 1220, // 747.5, /// 762.5, /K905; femur breadths—/*/ 
22.5, / 1 10, // 100, /// 92.5, IV 87.5. Opisthosoma 
contains large, granular horseshoe-shaped organ (open 
end at posterior), possibly gonad. Spermapositor short, 
breadth about half that of one genital shield, setae dg 
about half length of JZg. 

Material examined: Holotype male (N198317), seven 
paratype males (N198318-N198324), three paratype 
females (N198325-N198327), litter and sparse moss, 


REC. S. ALJST. MUS. 19 (4): 39-67 

Mu\\ 19H5 


2z 2J 

3Sa/ / ^ ^ 
3JZa J6 






,. chl 

FIGS. 13-16. Cmlonittjcilwrmente n.sp., malt cxcepi when utlicfwfec indicated, 13, noimn, 14, (unale idiosiernurn; 15, gnathostcrmini; 16. 
rifchl chelicera. anlcrior bUifaee. 







ad2 Pd2 /Sol 






ITCiS. 17-19. Crototiia jethurmenie n.sp., male; 17, leg I, dorsal surfaee of genu, tibia and tarsus; 18, leg tarsi, posterioi surface of distal 
ends; 19, leg III. posterior surface of trochanter and femur. 


Kl t . s \l st, Ml.s. m ( 4): $Wi 

,V/<;i, /W.I 

under FMcalyptus oblique Ml Lofty, 9.5.1974, D. C. 

Distribution: South Australia — Aa; Mr. Lofty, 
sclcrophyll open-forest, II i5/8). 

Remarks: Because one male lias little pigment or 
cerotcgumen! and therefore is easy to examine, it is 
designated holotype in contrast to the usual procedure 
of so designating a female. 

Luxton (1982) divides Cmionia into live species- 
complexes, amongst which C. jethurmerue would be 
grouped in Che eophinaria-complex. The inclusion of C. 
jethurmerae requires modification of the characteristics 
of that complex in thai hysteronota! setae 71 and SI are 
approximately equal in length, also setae j\ is l horn- 
like and straight. The curved lateral flap on ihc 
apophysis to seta -I appears unique within the genus. 


Nanhermanniidae Sellnick 1928: 17 
Nanhermanniidae: Woolley and Higgtris, 1956: 9i& 

Type-genus: Nanlwrmannia Berlese, 1914: 100. 

Diagnosis: Clinofissurac. Gnathostemal A-shaped 
mentocoxal fissure present Three pairs of adoral setae. 
Rostral tectum without median incision. Proteronoial 
plasmtc seta z2 baculiform or dilated and filiate disially. 
length tx-3x distance J2-z2. Hysteronotal seta./4 present. 
Mysteronotal gland absent. Notal setae z\ and 75 not 
on apophyses. Coxites //, Iff and IV may be 
hypertrichous. Setal file Sg includes two setae, combined 
setal file JZg on median margin of genital shield, 
tdiosoma almost covered in continuous shield except for 
relatively well separated genital and anal orifices and 
piehystcronotal fissure extending venttally (? anterior 
part of ventrolateral longitudinal fissure) as cresccntrie 
split nearly meeting midway between genital and anal 
orifices. Discrete preanal shield but internal under 
anterior margin of anal shields. Palp trochanter fused 
to femur, tarsus with seven setae. Tarsus / with one oi 
two pairs of dorsolateral supernumerary setae 
(solenidiuni5o1 level with them). Nymphs without Miiall 
shields around hysteronotal setal bases. 

Distribution: Probably cosmopolitan. Beside 
Nanhennannia, Masthermannia may be cosmopolitan. 
Cynhermannia is tropical (Cuba, NTa; Thailand, Os) 
as well as occurring in Japan (Pe). Whilst 
( >r,mohermannia is known from southern Japan (TV) 
and New Guinea (Am). 

Found in woodland and forest litter (both 
fermentation and humus layers). 

Remarky The only thoroughly described speeies in 
Nanhermanniidae is Masthermannia (as ftisthermanniu) 

nematophora (Grandjean, 1954b). The family was 
revised by Woolley and Higgins (1956). Although two 
genera and a number of species have been described 
since (ten, the form of Nanhermanniidae is without 
great variations from that oi the original speeies. 

Nanhermanniidae includes the following four genera: 
Cosmohermannia Aoki and Yoshida, 1970; 
Cynhermannia Balogh, 1958; Masthermannia Berlese, 
1914; Nanhennannia Berlese. 1914. 

V 4 NHERM4 NNI4 Berlese 

Nanhennannia Berlese, 1914: 100. Type designation 
(original): "Ilemtannia nana Nie"'. 

Type-species: Nanhennattnia nana (Nicolel, 1855; 458). 

Diagnosis: Nanhermanniidae. Hysteronotal setae not 
on swollen tubercles* simple, cither setose or lanceolate, 
sometimes minute forwardly directed proximal spur. 
Posterior margin of hvsteronotum evenly convex without 
prominent protuberances. Coxites ///, /l/\virh two to 
lour and three or four setae respectively. Trochanter IV 
wilh three dorsal setae 

Distribution: Probably cosmopolitan. Canada (Nn); 
Washington (No); Colorado, Idaho (Nr); Maryland, 
North Carolina (Na); Chile, Patagonia, Peru (NTc); 
Angola, Rhodesia (Ee); St. Helena (Es); Europe— most 
northern records Iceland and Kola Peninsula (Pe); Italy 
(Pm); Allay Mountains, kunashir Island, Sakhalin 
Island, Samarkand (Ps); Japan (Pc); Thailand (Os); 
Philippines (Om); New Guinea (Am); Queensland, 
South Australia (Aa); New Zealand (An). 

Remarks: Nanhennannia is the most widely recorded 
genus in the family and currently includes at least 
nineteen species. Balogh and Mahunka (1978), in 
describing N. domroW from Queensland, refer to a 
thaiensis-com\)\ex based on N. thaiensis Aoki, 1965b 
characterized by "medially confluent posterior 
protuberances of the prodorsum and that they do not 
have longitudinal fut rows among interlaineliar setae" 
for convenience, J will redefine the fhaeins/sxomp\c\ 
which Includes the species from this study. 


Diagnosis: Nanhermannia. Pair of posterior 
proteronotal protuberances basically semicircular, each 
with lour to seven minor protuberances. Furrow between 
setal pair,/2 absent or shallow, not breaking connection 
between protuberances, Genna and ubiae / and //seta 
n.»i bifurcate. 

Remarks Unfortunately the diagnosis lias to be based 
on the posterior sculpturing of the protcronolum, which 



is difficult to use for some intermediate species. The 
thuiensis-compkx is regarded as including the following 
m\ species: N. domrowi Balogh and Mahunka, 
1978— Queensland (Aa); N. Jorsslundi Karppinen, 
1958— Finland (Fc); N, gorodkovi Simikova, 
1975— Allay Mountains and Kunashir Island (Ps); N. 
grandjeani n.sp. — South Australia (Aa); H pectinata 
Strenzkc, 1953— Germany (Pe); N, thaiensis Aoki, 
1965b— rhailand (Os). 

Nanhermunnia grand jeani n.sp. 

(Tigs 20-26) 


General appearance and measurements: Light brown 
generally, darker around leg acetabula and posterior 
proreronotal protuberances. Shallow, clear 
ccToiegument, some adhering detritus around leg bases. 
5etae v claws, external malae, cheliceral digits clear and 
rcfraclile. Coarse puncta over much of soma excluding 
lateral regions of proteronotum, coxites genital and anal 
shields, and crescent shape on both sides of seta! file 
Sa (see Fig. 21). Much smaller bui deeper puncta located 
dorsally on proteronotum and central part o\' each 
coxite. Idiosomal length 637.5 (1); appendage lengths— 
ch 42.5, pa 55, / 265, // 250, /// 220, /! 280; femur 
breadths-/*/ 12.5, / 52.5, // 55, /// 40, /r 40. 

Pmsternwn: External malae without adaxial hyaline 
flap, but midanterior flap ventral to discrete tubercle, 
two rows of long cilia on dorsal surface. Three adoral 
setae, ao\ very fine. Coxites merged with each other and 
surrounding shields, although delineated by grooves. 

Proteronotum: Seta z2 club-like, ciliate on dilated area. 
Scia j2 has inconspicuous posterior spur at base. 
Middorsal T-shaped flat-topped mound with setae z\ 
at anierior end and setae z2 at end of lateral arms. 
Region often used in species diagnosis illustrated in 
detail (Fig, 24), Furrow between setae ,/2 shallow, 
accentuated by absence of small puncta that cover "P 
shaped mound. Posterior proteronotal protuberances 
obscure matching but smaller ventral protuberances (Fig. 

Opisthosternum: Shields distributed in manner unique 
to Nanhermannia. File JXg with nine .setae and Si? wirh 
three setae. Pores Zqfmd Saf present, other two pores 
illustrated (Fig. 22) regarded as hysieronorial (/;/). 

flysieronotum: Some setae (at least J2, Z2, 73. SI. 
S3, S5) with inconspicuous anierior spur at base. All 
setae blade-like, wirh hyaline dorsal and ventral flaps 
along entire length, not long enough in first three ranks 
to reach base of following seta. 

Append$$e& Chelicerae relatively small. On fixed digh 
five teeth including distal point, two small adaxial teeth 
parallel to main two proximal rceth. Movable digit with 

three teeth, including paired distal points and large 
proximal tooth. Setae: ch (2), pa (1-0-2-7), / (1-5-5-6-23), 
II (1-7-5-5-22), /// (5-2-2-3-18), /*' (1-3-2-2-16). Solenidia. 
pa (0-0-1), / (1-1-2), // (1-1-2), /// (1-L0), (V (1-10). 
Pretarsus with one claw. Terminal pair of plasmic setae 
on palp tarsus spine-like, in recess, Anlcrodorsal edge 
of tarsal Tibia with rcfraclile spur. Solenidia baculiforni, 
relatively short, never as long as associated setae. Tarsi 
/and /Teach with two supernumerary setae; possibly 
both dp. On tibiae ///and IV (only illustrated on IV— 
Fig. 25) seta &v conspicuously longer than scgnieni 

Soma! inclusio/is: No eggs. Ovipositor involuted, 
extended would be aboui 2s length of genital shield. 
Breadlh slightly less than thai of one genual sbirU 
Bears eight pairs o\' setae, two d% pairs about 0.6x length 
of setae J2g and 4\ length of setae pg. One small bolus. 
granular, particles many shapes, no complete cell walls 


Material examined: Holotypc female (N 198329), litter 
and sparse moss, under Eucalyptus ohltqiiu> Mt. Lofty, 
9.5,1974, D, C. Lee. 

Distribution: South Australia— Aa: ML Loftv, 
sclerophyll open-forest, 1 (1/8). 

Remarks: Amongst the //n//e>/s/.v-coniplex, differences 
between species in the description ol proteronotal 
protuberances and the intermediate furrow may be 
actual or represent different interpretations by authors. 
Within such variations N. %randjeani lies between N. 
pectinata and N domnwi. The shortness of the 
hysteronotal setae distinguishes N. gtvndjeam from N. 
domrowi, /Y fonslundi or N. gorodkovi N. pectinaia 
has no large puncta on the proteronotal T-shaped 
mound and seta zl is not dilated disrally. N. thaiensis 
has shorter, laterally biased protuberances, hysteronotal 
puncta are larger and uneven in size and shape whilst 
opisthosternal file Sa includes only two setae and the 
coxites bear only nine setae vent rally (3-1-2-3). 

Family HFRMANMIDAK Sellnick 

Hermanniidae Sellnick, J 028: 18. 
Hermanniidae: Woas, 1981: 7. 

Type-genus: Ilermannia Nicolet, 1855. 

Diagnosis: Clinofissurae. Gnathosternal A-shapcd 
mentoeoxal fissure present, at least median part. Three 
pairs of adoral setae. Rostral tectum without median 
incision. Proteronotal plasmic seta (zl) filamentous or 
club-like, length 0.75x4. 5.x distance Jl- "2. Hysteronotal 
seta 74 present. Hysteronotal gland present. Notal setae 
-1 and J5 not on apophyses. Coxites ///and IV setae 


R£C- S. AUST MUS. 19 (4): 39-67 

Ma\\ 1985 





FIGS. 20-24. Nanhermannia grandjeani n.sp., female; 20, notum and right pleura; 21, idiosternum; 22, gnathosternum; 23, right chelicera, 
anterior surface; 24, proteronotum, posterior protuberances. 


hypertrichous (3-1-4 or 5-5 or 7). Setal file Sg includes 
two to five setae, setal files Jg, Zg separate but may be 
close together. Idiosternum almost covered in 
continuous shield but well separated from notal shield. 
Discrete preanal shield, narrow (width about distance 
JZai-JZa2). Palp tarsus with nine setae. Tarsus /without 
dorsolateral supernumerary setae. Nymphs without 
small shields around hysteronotal setal bases. 

Distribution: Possibly cosmopolitan. Phyllhermannia 
found mainly in southern hemisphere (see below) whilst 
Hermannia confined to Palaearctic and Nearctic regions. 

Species of Hermannia occur on bark of living trees, 
regularly in forest litter, often at high altitudes, also in 
littoral habitats and salt marsh (Krantz, 1978). 

Remarks: A recent study of the Hermanniidae by 
Waos (1981) makes Phyllhermannia a junior synonym 
of Hermannia, so that ail species are in a single genus, 
with both types in the gibbia/convexa-complex. His 
study is disadvantaged by the brief descriptions of 
southern hemisphere species, i.e. those previously 
grouped in Phyllhermannia, The below r description is the 
first comprehensive study of a Phyllhermannia species, 






FIGS. 25-26. Nanhermanniu ^nmdjeam n.sp., female; 25, legs, dorsal and posterior surfaces of genua, tibiae and (arsi; 2fi, leg III posterior 
surface of trochanter and femur. 


kK\ S. AUSI, MUS. IV [4fc 3'if'? 

A An. wj; 

and character states such as ihe reduced sclaiion of the 
palp coxire, palp femur and legs, as well as the shape 
of adoral setatfoZ, suggest that PhyUhermannia may still 
be a valid taxon. Therefore. 1 have chosen what may 
appear a weak character (the position and shape of seta 
;;!) to diagnose (he two genera, because it has always 
been described and it makes a conservative grouping, 
requiring only one species to be excluded from 
PhyUhermannia, This is a temporary measure until 
PhyUhermannia phyllophora is properly described. 

The following two genera are included in 
Herman nndae: Hermannia Nicolet, 3855; 
PhyUhermannia Berlese 1917a. 


PhyUhermannia Berlese, 1917a* 65. Type designation 

(original): "Hermannia phyllophora Mich! 1 
PhyUhermannia: Tragardh, 1931b: 576. 
Hermannia (in part) Woas, 1981- 36. 

Type-species. PhyUhermannia phyllophora (Michael 
1908: 140). 

Diagnosis: Hermanniidae. Proteronotal seta z\ with 
distal half tapered off to a point, and marginal, lateral 
to line v /l-/2. Opisthoventral shield with transverse strip 
between genital and anal shields. Mentocoxal fissure 
usually complete, reaching edge of gnaihostcrnurn 
(exception: PhyUhermannia tuberculata). 

Distribution: Widespread in southern hemisphere, 
known range extending into northern hemisphere along 
western border of Pacific ocean. Chile, Juan Fernandez 
Islands (NTc); Cape Province, NataJ (Es); Madagascar, 
Mauritius (Em), Tanganyika (Ee); southern Japan (Pc); 
Thailand, Vietnam (Os); Java, Philippines (Om); South 
Australia (Aa); New Zealand (An); Pumas Arenas (Sm). 

Remark*: PhyUhermannia was established without any 
diagnosis, probably on The basis of the leaf-like leg setae. 
The first detailed consideration of the genus was by 
Tragardh (19.11b) and keys to some species are given by 
Aoki (l965b) and Balogh and Mahunka (1966), Woas 
(1981) regards this genus as a synonym of Hermannia 
and includes most species in his gibha/convexa<omp\ex. 

The diagnosis of the genus used here is weak since 
the character states of the opisthovcntral shield and 
mentocoxal fissure also occur in the small (after the 
exclusion of PhyUhermannia species) gibba/ 
convexa<omp]e\ within Hermannia. My approach has 
been to maintain PhyUhermannia until more extensive 
descriptions of included species, especially the type, are 
available. Only one species has to be excluded from this 
genus: Hermannia aerolata (Aoki, 1970) from Japan. 
This is still grouped in th? gibbia/convexa-compkx as 
by Woas (1981: 36), 

Twenty-four species and one subspecies arc included 
in PhyUhermannia. One species (R africana Balogh, 
1958) has not been considered because of the insufficient 
description. Of the remainder, ten species are considered 
similar to the new species described below and these are 
grouped in a species-complex. 


Diagnosis: PhyUhermannia, Hysteronotal seta Z\ 
central, not migrated laterally in front of 51, distance 
Z\-J2 subequal to or less than J\-J2. Hysleronotal setae 
short, ./I not reaching J2 base. Apodcmcs between 
coxites fJI and //-/// oblique, at least at 22.5 ° angle to 
transverse axis. Genital setal file Zg with at least one 
seta twice length of Sg setae, or more. Leg setae never 
leaf-like or spatulatc and number reduced, tarsus I wiih 
23 or fewer setae and one solcnidium. 

Remarks: Members of the euseiosa-complex arc 
usually only known by idiosoma! character states, I he 
three species from Mauritius, with only one setae in 
opisthosicrnul file Sg. may form a separate complex, P. 
tuberculata from Chile is included although unique 
within the genus in having a restricted mentocoxal 
fissure as in many Hermannia species. 

Eleven species arc included in the complex. R 
bimaculafa Hammer, 1979 — Java (Om); P. eusetosa 
n.sp.— Sout h Austral ia ( Aa ); P folia ta H amrner, 
J 966— New Zealand (An); P. mauritii Mahunka, 
1978 — Mauritius (Em); R modesia Mahunka, 
1978 — Mauritius (Em); P mollis Hammer, 1966 — New- 
Zealand (An); P. pacifica Hammer, 1972— Tahiti (Ap); 
/? pauiiani Balogh, 1962— Madagascar (Em); P rubra 
Hammer, 1966— New Zealand (An); P tremictu 
Mahunka, 1978— Mauritius (Em); P. tuberculata 
Covarrubias, 1967— Chile (NTc). 

PhyUhermannia eusetosa asp. 
(Figs 27-32) 

General appearance and measurements: Red -brown, 
covered in ccrotcgument, thin with sparse adhering 
dettitus. Beside small tubercles laterally and around 
coxites, acetabula and genital and anal orifices, soma 
(excepting proteronotum) covered in low, minute, 
superficial bumps, with pale strips between forming a 
reticulate pattern. All shields covered with fine puncta. 
Similar puncta and pattern on legs. Idiosomal length 
795 (25, 675-947.5); appendage lengths (for holotype, 
915)— ch 57.5, pa 90, / 580, // 430, III 440, TV 585; 
femur breadths— pa 27.5, HI7.5, U 105, III 90, /K95. 

Pmuernum; On external malae median hyaline Hap 
horizontal with posterior notch through which adoral 
seta ao3 protrudes. Internal mala with distal spike 
leading back lo minute dorsal furrow, Lateral refraclile 



part of external malae bears two rows of cilia on dorsal 
surface. Lateral and central gnathosternum with finely 
punctate patches as illustrated (Fig. 29). Coxites all fused 
into one shield but clearly delineated by grooves, deep 
along midsternal line. 

Rroteronotwn: Seta z2 vermiculate, but slightly dilated 
and ciliate distally. Bothridial cavity for seta z2 with 
number of short pockets and one long pocket. Two pairs 

of sharp tubercles near posterior margin face backwards, 
dark and conspicuous although small: one lies at 
posterior end of ridge running backwards from 
acetabulum /, other lies posterior to setae j'2 and z2 and 
equidistant from both. 

Opisthosternum: Continuous opisthoventral shield 
with strip between genital and anal shields, broadly 
fused to coxites, but not fused together posterior to anal 





— N. 


FIGS. 27-30. PJiylfheivuttwia eusetosu n.sp., female; 27, nolum; 28. idioslcrnum; 29, gnathosternum: 30. left ehelieera, anterior surface. 


FtEC S. AUST. MUS. 19 (4): 39-67 

May, 1985 





FIGS, 31-32. Phyilhermanma eusetosa n.sp., female; 31, legs, dorsal and posterior surfaces of genua, tibiae and tarsi; 32, leg 111 posterior 
surface of trochanter and femur. 


shields. Groove behind cox he IV. Cresent-sbaped 
thickening of integument behind acetabulum /I K Anal 
Shield wilh longitudinal ridge ending anteriorly in 
tubercle that fits around preanal shield. Both this 
tubercle and crescent-shaped thickening behind 
acetabulum IV conspicuously dark-coloured, although 
with pale shading (Fig. 28) because I hey stand proud 
from surrounding shields'. Pore Sez/present but not Zxtf 

llysteronoium: Two pairs of tubercles near anterior 
margin, ventral to eireumhysteronoial fissure and 
apposing protcronotal tubercles, central pair small and 
inconspicuous. Pair of dark semicircular integument 
thickenings behind first rank of hysieronotal setae (./I, 
71. SI), from which central arm borders on 
inconspicuous furrow running back to bend as faint 
ridge around seta ./4. All hysieronotal setae blade-like. 

Appendages: Chelieerae relatively small. Both digits 
with four teeth, terminating in paired, neatly parallel 
teeth (including distal points). Palp femur with 
ameroventral flange. Secae: eh (2), pa (0-1-1-3-9), / 
(0-6-5-5-22). // (1-7-5-5-17), /// (2-3-2-4-15), IV 
(2-4-3-4-14). Solenidia: pa (0-0-1), / (1-2-1), // (III), 111 
(1-1-0), IV (0-1-0). Pretarsus with single claw bearing pair 
of inconspicuous dorsal cilia files. Terminal pair of 
plasmic setae on palp tarsus spine-like, with anterodorsal 
flange around their bases. All but one solendiutn 
baeuliform, relatively shorty less than 0.5\ length of 
segment bearing them. Solendium sol on tibia 1 
flagclliforrru vctv long, about equal to length of genu, 
tibia and tarsus I together. On genua /, II, III and on 
all tibia (excluding so\ on tibia !) solenidia coupled with 
dorsal setae, whilst solenidia on tarsi A II and .vol on 
tibia I at most only associated with dorsal setae. 
Reticulate pattern on dorsal and lateral surfaces of most 
leg segments (Fig. 32 — p3rt drawn on femur), but not 
on dorsal surface of trochanter or any part of tarsi. This 
pattern consists of daiker raised ridges unlike similar 
hysteronotal pattern. 

Soma! inclusion^ Amongst twenty-five registered 
specimens, three contain two eggs, one contains three 
eggs, five contain four eggs, remainder without eggs. 
Eggs about 240 (225-265) long, ellipsoid, with uniform 
smooth surface, Ovipositor 2x length of genital shield. 
Breadth about l.5x that of one genital shield. Bears eight 
pairs of setae, two tig pairs subequal in length to setae 
Z#2 but thorn-like (x2 breadth) and more retractile, three 
pg pairs about length of genital shield away from three 
fffcg pairs. Only one bolus seen wilh mainly 
unrecognisable fragments, but some spherical spores and 
one multicellular strip of tissue, 



Material examined; Holotype female (N 198330), and 
twenty-four paratype females (NI983417-NIW3440), 

under Fiucalyptus obliqua, sclerpphyll forest, Mt Loft>; 
9 5.1974. 

Distribution: South Australia— Aa: Ml. Lofty, 
sclcrophyll forest, 118 (6/8), 

Remarks: P. euselosa is distinguishable from other 
members of the eu5<?/avtf-complex in possessing tbe 
following combination of character states; hysteronotal 
setae blade-like without cilia, no proteronotal ridge i tins 
either between setae jl-jl or seta jl and the median 
tubercle near posterior margin; coxite /Shears five setae. 


I am indebted to the Interim Council of the Australian 
Biological Resources Study for funds for equipment, 
and to the Science and industry Endowment Fund for 
funds for travel in connection with this project. 

Special thanks arc due to Mr D. MacParlanc. 
Commonwealth Institute of Entomology, London, for 
an English translation of a paper written iu French 
(Grandjean, 1954a) and to Dr G. Wauthy, Catholic 
University of Louvain, Belgium, for a gift of specimens 
of Eulohmannia rihagai. Thanks are also due to Dr 
B, C M. Jamieson, University oi Queensland, Brisbane 
and Dr R. A. Norton, S.U.N.Y. College of Environ- 
mental Science and Forestry, Syracuse, for commenting 
on the manuscript 

My greatest debt of gratitude is to Ms Jenni Thurmer 
for her excellent illustrations and to Mrs Debbie Melloy 
for typing this manuscript. 


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ULHAN. V. M. 1978 Ctttnk$H iabradonf.a (Aearil Onhjiei. 

Canu.siidac): a new soil initc- speeds ffOrtl UK Can&dlSil 

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Eu-Rl USE, A. J9l7a. Cemuria prima di Acan nuovt. «tv//u XII: I9-»V7. 
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BFRI ESE. A. 1017c Ceuluria tcr/a Ji Acari nuovi Rcdto 12: 2693% 
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by Wolfgang Zeidler 


Type-specimens of 21 species of Conus in the South Australian Museum are catalogued and 
illustrated. All are holotypes or syntypes. Of these at least three, C. cumingii Reeve, 1848, C. 
metcalfei Angas, 1877 and C. Sydney ensis Sowerby, 1887 are almost certainly not types while the 
status of another two, C. kermadecensis Iredale, 1912 and C. rossiteri Brazier, 1870 is in doubt. 
Only 10 of the specimens are from Australian waters, another 10 are from various South Pacific 
islands and one is from Mauritius. Species are listed alphabetically according to the original name 
of the species or variety. In addition, four species of cones recorded as types for which no reference 
can be found are listed in an Appendix. 




South Australian Museum, North Terrace, Adelaide, South Australia 5000 
(Manuscript accepted 29 August J 983) 


ZF.IDLER, W, 19H5. Mutlusc typc-spocimens in the South Australian 
Museum. 2. Gastaipoda: Conidae. Ret. S. Atisf, Mu\. I9<5): 69-75 

"type-specimens of 21 species of Conus in the South 
Australian Museum are catalogued and illustrated. All 
are holotypes or syntypes. Of these at least three, C. 
cumingii Reeve, 1848, C metcalfei Angas, 1877 and C 
sydneyensis Sowerby, 1887 are almost certainly not types 
while the status of another two, C kermadecensis 
Iredale, 1912 and C rossiteri Brazier, 1870 is in doubt. 
Only 10 of the specimens are from Australian waters, 
another 10 are from various South Pacific islands and 
one is from Mauritius. Species are listed alphabetically 
according to the original name of the species or variety. 
In addition, four species of cones recorded as types for 
which no reference can be found are listed in an 


Most of the indigenous Conidae collection in the 
South Australian Museum (SAM) was collected, or 
otherwise obtained by Sir Joseph C. Verco around the 
turn of the century. However, most of the exotic species 
were acquired by purchasing other people's collections. 
One of the most significant of these was the A. F. 
Kenyon collection which the Museum purchased early 
this century. Mrs Kenyon accumulated a considerable 
collection and acquired many types, particularly of 
cowries and cones. She also acquired much of the J. 
W. Brazier collection which accounts for about halt of 
the cone types listed here and only three of Brazier's 
species are not represented: C. cooki Brazier, 1870 (two 
syntypes in Australian Museum, Sydney), C. coxeni 
Brazier, 1875 (type in Coxen collection— lost?) and C. 
sophiae Brazier, 1875 (type in Australian Museum, 
Sydney). Unfortunately, several of the original labels are 
missing and the type status of some specimens is in 
doubt. There are also a number of specimens which have 
been labelled "type** but for which no reference can be 
found and these are listed in the Appendix and are only 
included because future workers may stumble upon a 
reference to them. 

The South Australian Museum holds 21 Conus 
"types" of which three, C cumingii Reeve, 1848, C 
metcalfei Angas, 1877 and C sydneyensis Sowerby, 1887 
are almost certainly not types while the status of another 

two, C. kermadecensis Iredale, 1912 and C. rossiteri 
Brazier, 1870 is in doubt. 

The species are listed under the original name at the 
time of description and in alphabetical order of species 
or variety. All of the types arc represented by shells only, 
and only two have the operculum with them indicating 
that they were alive when collected. Measurements for 
height and maximum width are given for each specimen, 
as these were often inaccurate or not given in the original 
description. Each specimen has also been photographed 
(Figs. 1-3) as the types were rarely figured and 
photographs of only a few have appeared in recent 

The present status of each species is, in most cases, 
according to Wails (1978). 


Conus kenyonae var. arrowsmithenste Brazier, 1896 

(Fig. la) 
Proc linn. Soc. New South Wales 21. 346. 
Holotypc: SAM Reg. No. D5459. Kenyon No. 337. 
Dimensions, 35.5 X 20.5 mm (Brazier gives 36 x 21 

Type-locality: Arrowsmith Island, Marshall Islands; 
collected by J. Brazier, 22.ix.I872. 
Present Status: Synonym of C distans Hwass, 1792, 
Remarks: A very worn specimen with Brazier's original 
label. Considered a juvenile of C. distans. Type figured 
by Cotton (1945), pi. 4, Fig. 4. 

Conus barbara Brazier, 1898 

(Fig. lb) 
Proc. Linn Soc, New South Wales 22; 78-1. 
Holotypc: SAM Reg. No. D6176. Kenyon No. S3. 
Dimensions, 40.0 x 20.3 mm (Brazier gives 40 X 
20 mm). 

Type-locality: Solomon Islands; collected by Mrs A F. 
Kenyon, date of collection unknown. 
Present Status: Synonym of C monachus Linnaeus, 

Remarks: A worn specimen with Brazier's original label. 
Type figured by Cotton (1945), pi. 4, Fig. 2. 


Kl l . S. Al'ST Mis )<) (5) fly.75 

AA/r. /ItfW 

Conus cumingii Reeve, 1848 

(Fig. Ic) 
Conchologia Jconica 1 (Conus Suppl.): PI. 3. species 282. 
Holotype: SAM Reg. No. D6208. Kenyon No. 372. 
Dimensions, 48.5 x 24.5 mm (Reeve gives none). 
Type-locality: None given on any labels but Island of 
Mindanao, Philippines; collected by Mr Cuming, 
according to Reeve. 
Present Status: Valid species. 

Remarks: This specimen cannot be Reeve's type oi* C 
cumingii despite the label which says "Reeve's lype". It 
does not agree with Reeve's description and is too large 
for the species. It seems to be C. anemone Lamarck, 
1810 and was not listed by Cotton (1945). 

Conus flinders'! Brazier, 1S98 

(Fig. Id) 
Proc. Linn. Sac. New South Hales 22: 780. 
Holotype: SAM Reg No, D14129. Kenyon No. 139. 
Dimensions, 28.0 x 15.0 mm (Brazier gives 29 x 
19 mm). 

Type-locality: Flinders, Western Port, Victoria, under 
stone at low water. (Not Flinders Island, Bass Strait). 
Probably collected by Mrs A. F, Kenyon, date of 
collection unknown. 

Present Status: Synonym of C anemone Lamarck, 1810. 
Remarks: This specimen does not match up with the 
dimensions given by Brazier, bul as (he original label 
by Brazier is present and in the absence of any other 
possible types, we must accept it as Brazier's type and 
conclude that his measurements are inaccurate. Cotton 
(1945) maintained that he had not seen the type. 

Conus frostiana Brazier, 1898 

(Fig. le) 
Proa Linn. Soc. New South Wales 22: 781. 
Holotype: SAM Reg. No. D6170. Kenyon No. 138. 
Dimensions, 18.2 x 9.5 mm (Brazier gives 18 > 
10 mm), 

Type-locality: Solomon Islands; collected by Mrs A. F 
Kenyon, date of collection unknown. 
Present Status: Uncertain. Walls (1978) thinks this 
species could be synonymous with C. nigmpunctutus 
Sowerby 1858 or C. monachus Linne, 1758 or even C 
magus Linne, 1758. I am inclined 10 believe that Uie type 
may be a small G monachus. 
Remarks: Specimen with Brazier's original label. 

Conus kenyonae Brazier, I896 

(Fig. If) 
Proc. Linn. Soc. New South Mates 21: 346. 
Holotype: SAM Reg. No. D14194. Kenyon No. 336. 
Dimensions, 420 x 23.5 mm (Bra/.ier gives 43 • 
24 mm). 

lypc-localiiy: Shark's Bay, Western Australia; collected 
by Mr Podesta. date of collection unknown. 
Present Status: Synonym of C. distans Hwass, 1792. 
Remarks: A very worn specimen with Brazier's original 

label. Considered a juvenile of C distans. Type figured 
by pcrtfcfti (1945), pi. 4, Fig. 6. 

Conus kermadecensis Iredale, 1912 

(Fig. Ig) 
Ptoc. Mai Soc, Lund 1 1K 3 >: 227, pi. 9, Tigs. 15 and 16. 
Syntypc: SAM Reg. No. D(>1 69. Kenyon No. 420. 
Dimensions, 43 x 23.4 mm. 
Type-locality: Original label lost? Iredale gives Sunday 
(-Raoul) Island, Kermadec Group. 
Present Status: Valid species, 

Remarks: Specimen with operculum, corresponding 
generally to Iredale's description. The lype status of the 
specimen, however, is in doubt as Iredale staled that the 
type would go to the Canterbury Museum. New Zealand 
(which apparently has two) and the paratypes would go 
to the Australian Museum, Sydney. The Australian 
Museum has three lots of C. kermadecensis collected by 
Iredale and Olivet from the type-locality, but none have 
been designated type or paratypes. Since Iredale 
mentions only three specimens, and two are accounted 
for, the above could be a syntypc corresponding to the 
one Iredale measured as 42 X 25 mm. Cotton's (1945) 
measurement of 45 mm for this specimen is obviously 
an error. 

Rhi:,oamus kiemae Cotton, 1953 

(Fig. Ih) 
Trans. Roy. Soc. South Attst. 76: 24, pi. 3, Figs. I and 3. 
Hololypc: SAM Reg. No. DI4465. Dimensions: 46.5 * 
26.0 mm (Cotton gives 47 x 26 mm). 
lypc-localiiy: Corny Point, Yorke Peninsula, South 
Australia; collected bv Miss M Klem, date of collection 

Present Status: Conns kiemae (Cotton, 1953), valid 

Remarks: Most of the specimens mentioned by Cotton 
(1953) arc in the SAM collections but none were 
designated paratypes. A juvenile specimen (SAM Reg. 
\n_ 1)16233}. collected with the type, was also figured 
by Cotton (pi. 3, fig. 2), 

Conus metcalfei Angas, 1877 

dig. 2a) 
Proc. Zool. Soc. (London): 173, pi 26, fig 13. 
Holotype: SAM Reg No. D6207, Kenyon No. 200, 
Dimensions, 17.6 X 9.5 mm (Angas gives 21.2 \ 
12.7 mm). 

lype-locality; Dredged at "Sow and Pigs" reel". Port 
Jackson, New South Wales; probably collected by J. 
Brazier, date of collection unknown. 
Present Status: Synonym of C atigasi Tyron, 1883. 
Remarks: J his specimen is almost certainly not Ihe lype. 
The original label is missing and the measurements do 
nut correspond to those given by Angas. The type is 
most likely in the Australian Museum* Sydney who have 
a specimen (C103598) with an old label marked "Type" 
measuring 20.5 x 11.4 mm which is a closet 



FIG. 1. a. Conus kenyonae van arrowsmithensis, holotype 35.5 x 20.5 mm ( x 1): b. Conus barbara, holotype 40.0 x 20.3 mm ( x 1): c. Conus 
cumingii, 48.5 x 24.5 mm (xl): d. Conus ftinciersi, holotype 28.0 x 15.0 mm (xl.5): e. Conus frostiana, holotype 18.2 x 9.5 mm (x2): 
f. Conus kenyonae, holotype 42.0 x 23.5 mm (xl): g. Conus kermadecensis, ? syntype 43.0 x 23.4 mm (xl): h. Rhizoconus klemae> 
46.5 x 26.0 mm (xl). 

approximation to Angas's description, 
confused with C. metcalfii Reeve, 1843. 

Not to be 

Kenyonia pulcherrima Brazier, 1896 

(Fig. 2b) 
Proc. Linn. Soc. New South Wales 21: 347. 
Holotype: SAM Reg. No. D6181. Kenyon No. 175. 
Dimensions, 28.0 x 9.4 mm (Brazier gives 28 x 
10 mm). 

Type-locality: New Hebrides; collected by Mrs. A. F. 
Kenyon, date of collection unknown. 
Present Status: Uncertain. This is almost certainly not 

a cone. It is most likely a turrid similar to Conopleura 
Hinds, 1844 (Cernohorsky, 1974). 
Remarks: A worn specimen with Brazier's original label. 
Not to be confused with Conus pulcherrimus Brazier. 
1894 or Conus pulcherrimus Heilprin, 1879 — a fossil. 

Conus pulcherrimus Brazier, 1894 

(Fig. 2c) 
Proc. Linn. Soc. New South Wales 9(1): 187. 
Holotype: SAM Reg. No. D6172. Kenyon No. absent. 
Dimensions, 80.0 x 27.2 mm (Brazier gives none) 
Type-locality: Tanna, New Hebrides, on beach after 


U\ ■< . S. ALIST MUS. W (5): &$-73 

Mm 1985 

FIG 2 a. Corns metculfei, V.6 x 9.5 mm (x2): b. Kenyonia pukhetrima, holoiypc 28.0 x 9.4mm(x 1.5): e. Conus pukhenimus, holotype 
80.0 x 27.2 mm (x 0.5): d. Conus tenia, lccloiypc 35.0 x 17.5 inm(xl); c. Conus rossiteri, ?holotype 10,6 x 6.4 mm (x3): f. Floraconus 
saundersi, holoiypc 57.0 x 32.0 mm (xl): g. Floraconus singleioni s holoiypc 43.5 x 22.0 mm (*1): h. Conus superstes, syntype 3.3 
x 1.7 mm <xl0). 

submarine volcanic eruption; collected in 1878; collector 

Present Status: Synonym of C. excelsus Sowerby, 1908. 
Remarks: A worn specimen with Brazier's original label 
indicating the species and locality but not the fact that 
it was Brazier's type. Cotton (1945) apparently ignored 
the original label and did not realize that Brazier had 
given a .short description under "Notes and exhibits" 
and thus described this specimen as the unique type of 
a new species, Asprella tannaensis. 

Not to be confused with Kenyonia pulcherhma 
Brazier, 1896 which is now considered a turrid or C 
pulcherrimus Heilprin, 1879 — a fossil. 

Conus remo Brazier, 1898 

(Fig. 2d) 
Proc, Linn. Soc, New South Wales 23; 271. 

Lectotype: SAM Reg. No. DI4128. Kenyon No. 186. 
Selected by Cotton (1945), p. 265, pi. 4, Fig. 9. 
Dimensions, 35.0 X 17.5 mm (Cotton gives none but 
Brazier gives 35 x 15 l /z-17 mm for the type). 
Type-locality: San Remo, Port Phillip Bay, Victoria; 
collected by Mrs A. F. Kenyon, date of collection 

Paralectotypes: SAM Reg. No. D15948, 8 specimens with 
same collection data as type. 

M0I II Si I Vl'l S(>li INK NS 2 ( .,.\SI ROW >i 'A. < (»M|»AI 

Present Stains; Synonym of C anemone Lamarck, 1810, 
Remarks: Specimens with Brazier's original label but the 
type had not been indicated. 

tonus rossiteri Bra/ier, 1870 

(Fig. 2c) 
Proc loot. Soc. (London) (1870); 109. 
Hololype: SAM Reg. No. D5975. Kenyon No. 312. 
Dimensions, 10.6 x 6.4 mm (Brazier gives approx. 16 
x 8 mm). 

Type-locality: Cape Solander, Botany Bay. New South 
Wales; collected by J. Brazier, date of collection 

Present Status: Synonym of C anemone Lamarck 1810. 
Remarks: This is a doubtful type-specimen as it is much 
smaller than the measurements given by Bra/ier, Also 
the original label is missing and there is no indication 
that this is the type apart from the museum register 
referring to it as "type" and Cotton's (1945, 1958) 
reference to it as the type. The Austalian Museum, 
Sydney, has a specimen registered as type in 1902, from 
material purchased from Brazier in 1889, measuring 
14.5 x 9 mm, 

Floraconus saimdersi Cotton, 1945 

(Fig, 2f) 
Rec, S. Aust. Mus. 8(2): 264, pi. 4, Fig. 8. 
Holotype: SAM Reg. No, D14198. Dimensions, 57.0 x 
32.0 mm (Cotton gives none). 
Typc-loeaTity: Levens Beach, Edithburgh, Yorke 
Peninsula, South Australia; collector and date of 
collection unknown. 

Present Status: Synonym of C. anemone Lamarck, 1810. 
Remarks: Two soecimens, SAM Reg. No. D16232, were 
with the holotype and may have been used in the 
original description but were not designated paratypes. 

I'loraconus singletoni Cotton, 1945 

(Fig. 2g) 
Rec. S, Aust. Mus, 8(2): 263, pi. 4, Fig. 10. 
Holotype: SAM Reg. No. D14195. Dimensions, 43.5 x 
22.0 mm (Cotton gives 45 x 22 mm), 
Type-locality: Western Port, Victoria; collected by C. J. 
Gabriel, date off collection unknown. 
Present Status: Synonym of C anemone Lamarck, 1810. 
Remarks: Considered an albino variant of C anemone. 
Two specimens, SAM Reg. No. D16236, were with the 
type but were not designated paratypes. 

Conus superstes Hedley, 1911 

(Fig. 2h) 
Z&ol Results F.I.S. Lndeavour 1904-10, part 1; 111, pi. 20, 
Figs. ^ 36. 

Synlypcs: SAM Reg. No. D15904, II specimens, the 
specimen figured here measures 3.3 x 1.7 mm. 

Type-locality: Forty miles south of Cape Wiles, South 

Australia in 100 fms; collected by F.l.S. Endeavour, 


Present Status: The species description is based on small 

juveniles, so (hat it would be difficult to determine the 

specific status until (he life histories of South Australian 

cones are belter known. 

Remarks: Specimens with the original label with the 

word ' k Co-type M and registration number "0.3800". 

Conus sydneyensis Sowerby, 1887 

(Fig. 3a) 
Thesaurus Conchvljorum, 5 (Conus SuppL): 260, pL 32 
(510), Fig. 694. 

Holotype: SAM Reg No. D6I83. Kenyon No. 192. 
Dimensions, 17.0 x 9.0 mm (Sowerby gives 24 
x 12 mm). 

Type-locality: Pbrt Jackson, New South Wales, collected 
by Brazier; date of collection unknown. 
Present Status: Synonym of C. aplustre Reeve, 1843 
according to Walls (1978) or of C angasi Tyron, 1883 
according to Cotton (1945). The specimen at hand looks 
more like C. angasi than C aplustre, 
Remarks: This specimen cannot be Sowcrby's type, even 
though the original label with the specimen, designating 
it as a type, appears to be in Brazier's handwriting. The 
locality data is given as Port Stephens and the 
measurements do not agree with the original description 
given by Sowerby. According to Moolenbcek (pers. 
commj the type appears to be in the Institut Royal des 
Sciences Naturelles de Belgique (Brussels). 

Asprella tannaensis Cotton, 1945 

(Fig- 2c) 
Rec. S. Aust. Mus, 8(2); 270. pi 4, Fig. 3. 
Holotype: SAM Reg. No. D6172. 
Remarks: Based on the same specimen as the type of 
C. pulcherrimus Brazier, 1894 also listed in this paper. 
The operculum figured with the type by Cotton (1945) 
probably belongs to another species as the type is worn 
and faded and was obviously collected devoid of the 

Hermes triggi Cotton, 1945 

(Fig. 3b) 
Rec. 5. Aust. Mus, 8(2): 267, pi. 4, Fig. II. 
Holotype: SAM Reg. No. D14324. Kenyon No.— absent. 
Dimensions, 61.5 x 27.4 mm (Coflon gives 60 
X 27 mm). 

Type-locality: New Hebrides; collector and date of 
collection unknown. 

Present Status: Synonym of C. ximenes Gray, 1839. 
Remarks: Described from a unique specimen from the 
Kenyon collection. 


U\ X ft ALSI. MLS. 19 (5): (W 75 

\am. ms 

FIG. 3. a. Conus sydneyensis, 17.0 X 9.0 mm (*2): b. Henne\ Triggh holotype 61.5 
x 16.0 mm |>:L5): d. Conus wotvesteri, holotype 47.6 / 24.5 mm <xl>- 

27.4 mm (xl): c. Conns waferhouseae, holotype 28.5 

Conus waterhouseae Brazier, 1896 

(Fig. 3c) 
Proc. Linn, Soc. New South Wales 21: 471. 
Holotype: SAM Reg. No. D5786. Kenyon No— absent. 
Dimensions, 28.5 X 16.0 mm (Brazier gives 30 
X 15 mm). 

Type-locality: Solomon Islands; collected by Mrs. G. .1. 
Waterhouse, date of collection unknown. 
Present Status: Synonym of C. distans Hwass, 1792. 
Remarks: Specimen with Brazier's original label. Type 
figured by Cotton (1945) pi. 4, Fig. 2. Another specimen 
(SAM Reg. No. D5787) with the type and marked "type 
of variety" was also figured by Cotton (pi. 4, Fig. 7). 
This specimen, from Mauritius, was introduced as a new 
variety by Kenyon (1906) who gave a brief description 
but it was not given a name as it was considered to be 
a juvenile of C. distans. 

Conus (Chelyconus) worcesteri Brazier, 1891 

(Fig- 3d) 
Proc. Linn. Soc. New South Wales 6: 276, pi. 19, Fig. 4. 
Holotype: SAM Reg. No. D6178. Kenyon No. 155. 
Dimensions, 47.6 x 24.5 mm (Brazier gives 48 X 25 

Type-locality: Island of Mauritius; collected by Mr 
Robiliard, date of collection unknown. 
Present Status: Synonym of C magus Linnaeus, 1758. 
Remarks: The original label by Brazier is missing but 
there is little doubt that this is the type. The specimen 
was collected live as the operculum is present. Type 
figured by Cotton (1945), pi. 4, Fig. 1. 


I wish lo thank Mr I. Loch of the Australian 
Museum, Sydney for supplying information on the cone 
types in that institution and Dr R. G. Moolenbeek, 
Instil ut Voor Taxonomische Zoologie, Zoologisch 
Museum, Universiteit van Amsterdam, The Netherlands, 
for his comments on the status of C kermadecensis and 
C sydneyensis. Thanks are also due to Mrs J. Forrest, 
Sourh Australian Museum, who took, and also 
prepared, the photographs. 


ANGAS, G. 1 . 1877. Descriptions of one genu* and twenty-five species 

of marine shells from New Souih Wales. Proc. Zoo/. Soc: 

(London): 171477, pi. 26. 
BRA/IER, J. IN70. Descriptions of three new species of marine shells 

from the Australian coast. Proc. Zoo!. Soc. (London): 109. 
BRAZIER, J. IK75a. Description of fourteen new species of shells 

fiom Australia and the Solomon Islands. Proc. Linn. Soc. New 

South Wales I: 1-9. 
BRA/ILR, J. 1875b. Descriptions oi' ten new species of shells, from 

ihc collection of Mr. Charles Coxen. of Brisbane, Queensland. 

Pmc. tool Soc- (London): 34-34, pi. 4. 
BRAZIER, J. 1891. Description of a new cone from Mauritius. Proc. 

JjntL StK. New Sourh Mules 6: 276, pi. 19. 
BRAZIER, J. 1894. Notes and exhibits. Proc Linn. Soc. New South 

Wales 9{\): 187. 
BRAZIER, J 1896a, A new genus and three new species of mollusca 

from New South Wales, New Hebrides and Western Ausiralia. 

Ptoc. Linn. Soc, New South Wales 21: 345-347. 
BRAZIER, J. 1896b. New species of cone from the Solomon Islands. 

Proc. Linn. Soc. New South Wales 21: 47t. 
BRA/IER, J. 189Ha. New marine shells from the Solomon Islands 

and Australia. Proc 1 frrft Soc. New South Mates 22: 779 782 f 
BRAZIER, .1. 1898b, Four new species ol Mollusca from Victoria. 

PMC Linn. Sot New South Mw/w r 23: 271-272. 
t ERNOHORSKY, W. IK74. The taxonomy of some Indo -Pacific 

Mollusca with description of a new species. Pari 2. flfce. Auckland 

Inst. Mus. II; 121-142, 



COTTON, B. C. 1945. A catalogue of the cone shells (Conidae) in 

the South Australian Museum. Rec. S. Aust. Mus. 8(2): 229-280, 

pi. 1-5. 
COTTON, B. C. 1953. New species and records of Mollusca from 

South Australia. Trans. Roy. Soc. South Aust. 76: 21-26, pis 2, 3. 
COTTON, B. C. 1958. Australian recent and tertiary species of the 

molluscan family Conidae. Publ. B. C. Cotton. 4 pp. Sept 1. 
HEDLEY, C. 1911. Report on the Mollusca obtained by the F.I.S. 

Endeavour chiefly off Cape Wiles, South Australia. Zool. Results 

FI.S. Endeavour 1904-10, pt. 1: 90-114, pis 17-20. 
IREDALE, T. 1912. New generic names and new species of marine 

Mollusca. Proc. Mai. Soc. (London): 10(3): 217-228, pi. 9. 
KENYON, A. F. 1906. Notes. Proc. Mai. Soc. (London): 7(1): 5. 
REEVE, L. A. 1848. Monograph of the genus Conus. Conchologia 

Iconica 1 (Conus Suppl.). 
SOWERBY, G. 1887. Thesaurus Conchyliorum 5 (Conus Suppl.). 
WALLS, J. G. 1978. Cone shells: a synopsis of the living Conidae, 1011 

pp. T.F.H. Publications. 


Specimens labelled or registered as "type" for which 
no reference can be found. 

Conus noumeensis Brazier, SAM Reg. No. D5729. 
Kenyon No. 69. (68.5 x 39.5 mm). 
Locality: Anse Vata, Noumea, New Caldonia. 
Remarks: Specimen with Brazier's label marked "Type 
specimen". Cotton refers to this specimen and says that 

it is C. chenui Crosse, 1857 and has nothing to do with 
C. suff usus var. Noumeensis Crosse, 1872 which is a 
synonym of C. marmoreus Linnaeus, 1758. 

Conus pitched Brazier, SAM Reg. No. D6171. Kenyon 
No. 247 (22.3 x 14.0 mm). 

Remarks: Specimen with Brazier's label marked "Type 
specimen" but no locality data. It is very similar to C. 
cyanostoma A. Adams, 1854. 

Conus sydneyensis var. sowerbyi Kenyon, SAM Reg. No. 
D6184. Kenyon No. 192 (27.0 x 15.5 mm). 
Locality: Port Stephens, New South Wales. 
Remarks: Specimen with Kenyon's label marked "type". 
Another specimen with a similar label but not marked 
"type" is also in the collection. The specimens seem to 
be juvenile C. purpurascens Sowerby, 1933. 

Conus wisemani Brazier, SAM Reg. No. D6179. Kenyon 
No. 199. (28.2 x 17.3 mm). 

Locality: Low Island, Trinity Bay, north Queensland. 
Remarks: Specimen with Brazier's label but not marked 
"type" yet registered as a type in Museum register. It 
seems to be a white form of C. suturatus Reeve, 1844. 




JUNE 1986 








by W. ZEIDLER and F. W. ROWE 



North Terrace, Adelaide 
South Australia 5000 


byE. G. Matthews 


Eight species and one subspecies of Brises Pascoe are recognized, keyed, and briefly described. 
Four of the species are new and named caraboides, katherinae, occidentalis and nullarboricus. The 
specific name duboulayi Bates becomes a subspecies of acuticornis Pascoe, and granulatus Carter 
becomes a synonym of the latter. There is a brief discussion of the cave and burrow frequenting 
habits of Brises. 

A number of former subfamilies and tribes of Australian Tenebrionidae are combined into the 
single tribe Cyphaleini and adult diagnostic characters for the latter are proposed. 





South Australian Museum, North Terrace, Adelaide, S.A. 5000 
(Manuscript accepted 5 November WH4) 


MATTHtW-S, K C\. 1986* A revision ol ilie troftophific germ* 9/vm 
['ascot, willi a discussion pf the Cyphaleini (Cok-opiera). Hec 
S Auu A*V* 19(6): 77 *)U. 

Tight species and one subspecies of Brises Pascoe are 
recognized, keyed, and briefly described. Four of the 
species are new and named earaboides, katherinae* 
occidentalism and nullarboricus. The specific name 
duboulayi Bates becomes a subspecies of acutkvmis 
Pascoe, and granulatus Carter becomes a synonym Of 
the latter. There is a brief discussion of the cave and 
burrow frequenling habits of Brises. 

A number of former subfamilies and tribes o! 
Australian Tenebrionidae are combined into the single 
tribe Cyphaleini and aduli diagnostic characters for the 
latter are proposed. 


The genus Brises Pascoe, 1869, comprises eight known 
species whose distributions are practically confined to 
the arid /.one of Australia, with most known records 
being within South Australia (Fig. 23). It is unique 
among Australian lencbrionidae in that the majority 
of species have been recorded from eaves and mammal 

Brises belongs to the very large and diverse 
Australasian endemic tnbe Cyphaleini (subfamily 
Tenebrioninae) which represents the most important ol 
the three dominant elements of the Australian 
lenebrionid fauna, the others being the Adeliini 
(subfamily Lagriinae) and Amarygmini (subfamily 
Tenebrioninae) (Kas/ab 1982). Having probably 
originated as forest-inhabiting, rotten-wood-feeding 
forms (as many still are) certain elements ol the 
Cyphaleini invaded the arid zone to become soil 
inhabiting as larvae and ground foraging as adults, but 
they do not remain active under daytime conditions as 
do many African desert tencbrionids. The concept of 
the tribe Cyphaleini adopted here largely conforms to 
that implied by Doyen and Tschinkel (1982) and includes 
all the members of the *5.u&tanwb'cs" Helaeinae, 
Nyckvoilinae, Briseinae, and Cyphaleinae of Carter 
(1926), a total of 46 genera and some 450 species in 
Australia and New Guinea, plus the 14 species oi' 
Munopeus Pascoe in New Zealand (Watt 1968), 

June, IWj 

The following abbreviations for institutional names 
are used in the text and follow the four-letter system 
proposed by Watt (1979): 

AMSA —Australian Museum, Sydney. 

ANIC — Australian National Insect Collection, 

BMNH— British Museum (Natural History). 
MVM A— Museum of Victoria (Natural History and 

Anthropology), Melbourne. 
QMBA —Queensland Museum, Brisbane. 
SAM A —South Australian Museum, Adelaide 
UQBA —University of Queensland I ntomology 

Department, St Lucia, 
WA MA— Western Australian Museum, Perth. 

The primary types of all the taxa of Brises have been 
seen by the author. 

The Limils of the Cyphaleini and Tenehrionini 

As discussed by Doyen and Tschinkel (1982), these two 
tribes belong to the "leuebrionine lineage" which is 
characterised in part by having the spermatheca derived 
from the original bursa copulatrix (which has 
disappeared as a recognizable entity), by the orientation 
of the eoxite and paraproct baculi of the ovipositor 
being transverse and oblique respectively (Fig. 1), and 
by (he defensive gland reservoirs opening between 
sternites 7 and 8 and being small and not reinforced by 
helical thickenings (Tschinkel and Doyen 1980). These 
three features are shared with (he closely related toxicine 
and opatrine lineages (best treated as further tribes of 
the Tenebrioninae) and sharply distinguish these lineages 
collectively (in effect the subfamily Tenebrioninae) from 
other subfamilies. The above authors do not propose 
any formal classification, but I equate the subfamily 
tenebrioninae with their tcnebrioninc, tovicine, and 
opatrine lineages. 

The Tenebrioninae (as understood here) are further 
characterised by a number of primitive features, 
including the almost complete absence of the clustered 
antennal sensoria termed "tenebrioid sensory organs*' 
by Medvedev (1977) and, inconsistently in apparently 
primitive elements, internally open fore coxal cavities 
and a subeubital fleck on the wings (Fig, 2). 

The two tribes Tenebrionini and Cyphaleini share all 
of the above features, at least in part, and in a natural 
system could not be satisfactorily separated. However, 


REC. S. AUSl. MUS. 19(6): 77-90 

June, W86 

the resulting group would be rather unwieldy on a 
world-wide basis and it is desirable for purely practical 
purposes to try to find an arbitrary line of separation. 
Watt (1974) suggests a larval character — a crenulate 
spiracular peritrcme, combined with a primitive adult 
character — the subcubital fleck, to characterise 
Cyphaleini. However, too few larvae are known for us 
to be able to understand the significance of the 
spiracular character, and the adult one is not only 
plesiomorphic but also by no means universal in the 
Cyphaleini. For practical reasons I prefer to draw the 
line at an adult apomorphic character — the absence of 
original elytra] striae — to recognise Cyphaleini, when 
combined with certain other features* 

Specifically, Cyphaleini may be considered to be 
members of the Tcnebrioninae which share the 
following adult characters (A— apomorphic; P— plesio- 
morphic): (I) absence of tenebrioid sensoria (P),(2) 
internally open fore coxal cavities (P), (3) the Ihird 
antennal segment longer than the preceding or 
following ones (A), (4) the original nine elytral striae 
entirely absent, the elytra cither non-striate or with 
secondary, usually supernumerary striae (A), (5) no 
sexual dimorphism in the shape of the fore legs (P), 
and (6) the paramcres of the aedeagus with a vestiture 
of minute spines (A?). As far as I know, all Cyphaleini 
share the above features, but conversely none of the 
features arc exclusive to the group. Meneristes Pascoe, 


accessory gland 

CuP and 1A 
parity merged 

FIGS, 1-4. 1- Ovipositor and internal female genital apparatus of Bri$es a. acuiiamus I'aseoe. 2. The tenebrionine wing venation {Mefwmtes 
australis Blcs.sig). Venation nomenelaiure after Medvedes (1968). 3. The eyplialeine wing venation (BrtM's a, iwuiicttrtus Pascoe). 4. 
The eoelometopine wing venation {Encyulesthus arravindis Maeleas >. 



in particular, is a iraiiMiioual genus between Cyphaleim 
and Tenebrionini, since it displays characters I, 2, 3, 
and 6, hul not 4 and 5, and ii was placed in the 
Cyphaleim by Wail. (1974), a posiUou supported by 
Doyen and 'Ischinkel (1982), I prefer to consider the 
elytral character (4) to be decisive, because of the ease 
with which ii- can be seen, and so place Meneristes in 
I he Tenebrionini. 

Additional characters which aa- frequent but not 
universal in the Cyphalcini are the presence 6[ a 
subcubital Heck on (he hind wings (P) (in ten of the 
20 wjnged genera examined, but not in Brisesh the last 
maxillary palpal segment triangular or securiform (A) 
(29 of 33 genera), a shortening of the paramercs in 
relation to the legmen of the aedeagus (A) (14 of 33 
eencra examined) and a sclerotisation oi the median 
lobe, often in a characlerislic form (arrow-shaped in 
ttrises) (A). In addition, eight of (he winged genera 
examined (Brise; Pascoc, Pierolit'/aens Brcmc, 
Emtephulas Kit by. Para phones Macleay, Ospu/us 
Pascoc, Baryhphia Pascoc, Anuusts Bates and 
Chartopteryx Wesiwood) display a venalional 
peculiarity illustrated in Pig. 3. whereby IA is broken 
at one spor and a now cross vein appears, usually only 
a- a stub, further proximad between CuP and I A 
(compare (he normal icncbrionine venal ion seen in 
Meneristes, Pig. 2). The latter configuration also 
appears jri at least 12 genera of Cyphaleim. so wing 
venation cannot be used reliably as a tribal character. 

The genus Tiiaenu Brichson clearly belongs in the 
Cyphalcini as here conceived since U shows all <he 
diagnostic features mentioned above except (3) and 
some of i he additional characters (a subcubital fleck, 
secuiifotm maxillary palpal segment, and sclerotisation 
of the median lobe). However, Jitaena is apparently 
also close to Ar/ystotw Bates of New Zealand and 
Calhsmila.x Bates of New Caledonia, all three placed 
in a tribe liiacnini by Kas/ab (I9K2). The latter two 
genera display the lull complement of original elytral 
striae (at least in some species) and so do not fall within 
the Cyphalcini as here arbitrarily defined. I have not 
closely examined Arfvsfona and Caitisnulax and prefer 
lo leave the question of their relationship (and therefoic 
that of litaenu) in abeyance. 

It should also be mentioned here that the subfamily 
Coelometopinae, which is sharply distinctive o\\ female 
gcnjiabe and gland reservoir characters (Doyen and 
Tsehinkcl 19S2), can often also be recognized by a 
eharacletistic wing venation in which the veins CuP 
and I A have completely merged for a short distance, 
wallowing erossvein Ciwi (Fig. 4). This modification 
is accompanied by a reduction of the vannal area. 
However, a lew eoelomet opines have a normal 
lenebrionine venation fWJ. Ieinwonomenes Chcvrohu. 
(hanotheca Pascoc). t oclomciopiuae strongly 
u -cubic lenebnonini in external form but can always 
be recognized without dissection by ihe piesencc of 

tenebrioid sensotia o\-\ the antennae, universal in the 
group. Ihe following coelometopine genera were 
erroneously placed in the Tcncbrioninae by Cartel 
(1926); llvpuulux Bates, ilyttissus Pascoc, hmyahsihas 
Motschulsky, Setenis Motschulsky, Promethis Pascoc, 
letru^onomenes Chevrolat, Oectosts Pascoe and 
Zophophilus Fairmaite (the correct placement of some 
of these genera has already been proposed by Doyen 
and Ischinkel (1982)) 

The Australian genera which I consider to be one 
lenebrionini are Asphahts Pascoc, Sloanea Carter, 
Monwisies Pascoc, an unnamed genus comprising the 
species colydioicles \ nehson, recrihasis Carter and 
.syc/neyanus Blackburn, all three erroneously pm In 
Menephilus, and Parataxicum Champion. 

The 46 genera o\ Cyphalcini will not be listed here 
but can be obtained from Carter's (1926) checklist 
Lindei the 'subfamilies" Hclaeinae, Nyeto/oilinae, 
Briscinae, and Cyphaleinac 

Suhyrmips of Cyphalcini and Recognition of times 
The placing of the cyphaleinc genera in four separate 
subfamilies by Carter (1926) suggests thai some natural 
gtoupings may exist within the tribe, perhaps al 
subtribal level. In the coune of Ihe present study I 
dissected representatives of 33 cyphaleinc genera but 
was unable to arrive al any arrangement, and believe 
that the former categories cannot be maintained at any 
level. There is no concordance in the pattern of 
variation of characters, whose slates eotvsetpiently 
follow a mo/aic disirihution. This is a characteristic 
feature of the Tenebrionidae as a whole and one which 
has so fat defeated all aUemprs to arrive at a convincing 
natural classification of the family (Doyen and 
Lawrence 1979; Doyen and Ischinkel 1982). 

Hrises itself was singled out by Carter in 1914, and 
placed in its own subfamily in 1926, because of a 
number of ostensibly unique leal arcs, specifically the 
narrowly triangular shape of the last maxillary palpal 
segment (normally strongly securiform in Cyphalcini), 
the narrow posterior intercoxal process, the elongate 
legs, and the long tibial spurs and tarsal claws. After 
a detailed examination of lepresentative Cyphaleini 1 
find it difficult to understand just what is so unusual 
about Bribes. All of ihe above characters recur in other 
genera (bur not in ihe same combination). I ike all 
cyphaleinc genera Briscs may be recognized only 
through a unique combination 0? £ number of 
characters, enumerated for hrises below under 
"Diagnosis 1 * It also has a distinctive habitus resembling 
that of cursorial carabid beetles, due mainly lo its 
relatively narrow prothorax and long legs. This charac- 
teristic build is the principal feature distinguishing ii 
from some species ol P/enfhelaens Hi erne, in which the 
omliues of rhe pmnotum and elytra arc confluent, but 
which in othet respects resemble Rrises. Pterohelaeus 
itself appears lo be a composite mvon. 


KEC, S. AUST MUS. 10(6): 77-90 

June, 1986 

Adaptations of Brises 

The Cyphaleini are a relatively primitive group of 
Tcnebrioninae which has radiated extensively in the 
Australian region in the absence of the subfamily 
Pimeliinae (Tentyriinae), which elsewhere tends to 
become dominant in similar \eric situations. Brises 
itself has evolved a strategy of utilising caves and 
burrows lor daytime shelter at Jeasi. Six of the nine 
taxa have been recorded front caves (caraboides, a. 
ocutieornis, a. duboulayu katheritwe, occidentalism and 
trachynotoides), and three of these (the two subspecies 
of acuticornis, and traehynotoides) from rabbit 
biJTrows as well, Two of the remainder are recorded 
from rabbit burrows only (blairi and parvicollis), only 
nuliarboricus being without any data on shelters used. 

G. B. Monteith {in tin.) writes that south of fiirdsvijlc 
blairi once came out at night in August in enormous 
numbers foraging on the crests of sandhills, even 
though it was freezing cold. ft a. acuticornis feeds 
inside caves and so, presumably, does kuthennae (sec 
accounts under species). Feeding habits of the other 
species are not known. Larvae which have been round 
in caves and burrows in association with Brises adults 
axe assumed to belong to the latter, but some Helaeus 
species also use burrows (Matthews 1985) and it is 
necessary to rear larvae through to confirm their 

By examination of gut contents and through 
observation of captive specimens of acuticornis I infer 
that the adults are genera) scavengers like most tene- 
brionids. In burrows they (and presumably all other 
species of the genus) are probably coprophagous on 
the droppings of the mammal inhabitants, and in caves 
they appear to feed on all faecal matter available and 
on carrion. 

Adaptation of Brises to life in caves and burrows is 
barely reflected in then structure. Long appendages, 
small eyes and a large hind body are features often seen 
in cavernicolous beetles, as is wing reduction, In all 
these respects Brises is only at the earliest .stage of 
modification, carabotdes being the most modified. 
Only two species appear to be flightless and only five 
have eyes measurably smaller than those of winged 
cpigean genera. Intensity of pigmentation is at normal 
levels, but paler specimens are common in duboulayi 
Altogether, the morphology of the species suggests that 
there is frequent migration from one burrow or cave 
to another. 


Brises Pascoe 

Bases Pascoe, 1869, p. 145; Carter, 1914, pp. 45, 46; 

Carter 1926, pp. 127, 145. 

Ephidomus Pascoe, 1869, p. 151; Carter, 1914, p. 45 


Type Species: Of Brises: Brises trachynotoides Pascoe, 
1869, by monotypy Of Ephidonius: Ephidonius 
acuticornis Pascoe, 1869, by monotypy. 

Description: Entirely black. 

Head: Anterior margin of clypcus concave. Clypeo- 
frontal suture complete or not. Basal membrane of 
labruni visible from above. Eyes entire but constricted 
by both canthus awl edge of occiput, the dorsal 
interocular distance 2I-2-5 times width of one eye, Gtilar 
sutures meet near middle, no gular pits, anterior edge 
of gula unmodified. Bridge of tentorium straight. Third 
antennal segment longer than 2nd or 4th. Tormae of 
labrum transverse, without prominent anterior 
extensions. Mandibles bidentate, inola not striate. 
Lacinia unarmed. Terminal segment of maxillary 
palpus narrowly triangular to sublinear (Fig. 5), of 
labial palpus subiinear. Angles of mentum anterior, 

Thorax- Margins of pronotum prominent but rarely 
strongly explanatc, anterior edge evenly arcuate. 
Outline of pronotum not confluent with that of elytra, 
narrower. Pronotum glabrous or with minute, very fine 
setae. No sharp prosternaJ keel. Fore coxal cavities open 
internally. Mesosternum strongly excavate to receive 
prosternal process, Metendosternite Y-shaped, with 
laminae. Scutellum visible, small, in form of 
subequilatcral triangle. Elytra glabrous or with minute 
setae, true striae absent, cannae present or not. 
Epipleura moderately wide, complete. Wings without 
subcubital fleck or with faint trace of one, with base 
of first branch of 1A broken and partial supplementary 
cross vein eu-a present (Fig. 3). Tarsi slender, eiaws 
long, equal in length to about : j o( last tarsal segment. 
Tarsal vestiture in form of long bristles. Tibial spurs 
equal in length to { A- { A length of metatarsus, legs 
slender, without carinae. 

Abdomen: liuercoxal process of first visible 
abdominal sternite narrowly triangular. Reservoirs of 
defensive glands small, simple. Ovipositor and internal 
female genital tract of tenebrioninc type (Fig 1) 
Parameres of aedeagus spinose, comprising */M of 
total aedeagal length, with backward prolongations, 
Median lobe arrow-shaped or simply expanded 
subupieally (Figs. 11-14). 

Total length 10-25 mm. 

Diagnosis: Eyes in dorsal view ovoid or subquadrate, 
not strongly transverse, iuterocular distance equal to 
2'/2-5 eye widths. Membrane at base of labrum exposed 
by concavity of clypcal margin. Last segment of 
maxillary palpus narrowly triangular. Pronotum 
glabrous, markedly narrower than clytral bases and not 
confluent in outline with clytral edges. Elytra glabrous 
or with minute setae, the lateral margins not expanded. 
Wings (when present) without distinct subcubital Heck. 
Legs slender, femora extending lor about W-W of their 
length beyond body sides. Tibial spurs equal to ] A to 



FIGS. 5-14. 5. Outline of terminal maxillary palpal segment, Brises trachynotoides Paseoe. 6. Antenna, Brises nuUarboncus n. sp. 7. 
Antenna, Brises a. duboulayi (Bates). 8. Hind tarsus, Brises caraboides n. sp., setae omitted. 9. Hind tarsus, Brises occidental n. sp. 
10. Hind tarsus, Brises trachynotoides Paseoe. 11. Aedeagus of Brises a. duboulayi (Bates) in side view. 12. Apex of aedeagus of Brises 
a, duboidavi (Bates) in ventral view, showing arrow-shaped end of median lobe. 13. Aedeagus of Brises irachynotoides Paseoe in side 
view, 14. Aedeagus of Brtses trachynotoides Paseoe in ventral view, median lobe also shown partly extracted. 


KM., S. AUSI MUS !•>«»>• 77-9(1 

//;//(•, /VAYt 

!/i length of metatarsus. Claws long, equal in length 
to about : < of last tarsal segment. larsi bristled 
beneath* not lomenlose. Intercoval proeess of first 
visible abdominal sternite narrowly triangular, 

Distribution (Fig. 23): All of South Australia except 
ihe southeastern portion; New South Wales west of the 
Great Dividing Range; southwestern Queensland; the 
Northern Territory from Katherine southward; the 
Nullarbor, Central, and North West districts of Western 
Australia. Probably occurring in all parts of Australia 
receiving less than 300 mm of annual rainfall, with 
populations substantially outside this area possibly 
isolated and restricted to eaves. 


1 {2) Humeral .mules strongly cxplanale. Llyiral length about 1.3 

lirnes basal elyrral s\ tcii h. Wingless (Kijj 241 

- - I hktirt (- aiiei 

Humeral antics rounded. Hyiral k-ntiih t.6 2,\ mnev h.ivil 
•d viral ■uidlb. Wimts present but may he reduced . , 2 

2 (0 Clural surkiev Willi punctures arranged in urns, scimeiinics 

inquilai. Granules Dr spines, il present, almoin ro\\> He;id 
and pronomm usuall\ shugieencd and puric-rale, lertrnnal 
anienintl Moment mure ov less acuminate (Pig. ?). No sexual 

dimoi phism ... 3 

tlynal surlaees coarselv reticulo -punctate and granulate, the 
punctures and granules never in rows on disL $\ least. Head 
and pronolurn densely vcrmiculatcot granulare. terminal 
anicnnal segment not uewnmaie, ai most somewhat 
narrowed apically (I ig 6). Mule with middle and hind 
Iciilorj modified (I i«s \9-22) ........ 8 

\ (2| Ptonotal surface shiny, TincK shajrrccned and punctate only, 

without granules 4 

PromMul surlaec malt, uiih at least a lew small I'ramtles 

4 Ol Hind body oval (Hg_ 28$ t lliyhilcss. Llvtra Willi itu intervals 
between low, rounded eartnae c.arseh and deuseU punc- 

latc, without granules , 2 caruhohics n. sp, 

Hind bod* oblona (I iv, 2n), wines normal- Hyiral lUfffce 
finely punctate, wnli or wiihout granules and earinac 

5 (4J Elvira without trace of ^pines, granules oi sharp canine 

3 acutfcnrms acuueortus (Pascoe) 
t.lyira with jy :» tew small ?.*ranutt".. may be dtstmeily 

tiraniilate and eannalc . , 

,4 ttvHthomis (Jubt.'tihtyi 05ates> 

6 tt\ Head and pronoimn eoaiscly HXttq&e and granulate, f'rovternal 

proves with the margins sharply raked bttVOC^B lore eo>je. 
I lyirjl shrlacc wdh short setae, more prominent laterally, 

elslral edges nearly straiahr in middle (I ie_ 29) 

......... 5 kathaitute n . sp. 

Head and pmuojum with small granules, the -air lace between 
i hem shagreencd and finely rtteose. PmMemal proofs Willi 
margin not raised, tlyira without setae, Uyiral ed±jes. 
dishiiaK com ex ihmmdioui ( t iyv. 10 ^nU 3!| 7 

7 f$J \ yes wider (liar. lon«, M/p.nalcd by about 4 eye Widths 

(lie P| llyiral caiinae wiy prurmtieni. ihe siulacc 

between them coIkjsc and disitncily granulate , . . 

- - - - - - h oi\'i<li't)uths tl. sp. 

Pvex about as lone as wide :uul separated by about 3 eve 
widths tJ ii_*. EBE}. llyiral catinae line, ihe mu late between rial, (iiiiiiiiely yranulalc. \vi|h fine supplementary 
longitudinal ndyes . , 1 paivivollls (UtackburtU 

8 (2) t lypeusin lower plane than Irons, dearly demarcated at base 
Kroitt etlgcsol\'ves strongly oblique(Pig. 15). Hctra WtiH ^ 
short humtral carina (I if ^2) S nttlit/t Intra us rt: ^p. 
C'lyix'us in same plane as Irons, noi sharply delimited at base. 
1 rota cdptt oi eyes more ov less transverse (I ig. \$). V iv pi 
withirin a humeral carina . , 9 tn/drnwioid^ Pascoe 

I. Brises htuiri Carter 
(Fig. 24) 

Brises Nam Carter, 1914, p. 58; Carter, 1926, p. 145. 

Description: Mentum wilh two pits and broad 
anterior membranous area. Terminal antennal segment 
apieally narrowed btn obtuse. Clypeus in lower plane 
than irons. Eyes with length subequal to width in dorsal 
view, separated by a distance equal to about 2Vi eye 
widths. Edges of prosternal proeess not raised. 
Pronoturn 1.7-1.9 times as wide as long with margins 
strongly c^planate. Pronolal surface aiutaeeous, with 
scattered very fine punctures. Flyiral surfaces with very 
small granules, denser anteriorly, and very small 
punctures more evident anteriorly and along sides, the 
surface between them aiutaeeous, without setae. Two 
incomplete raised eartnae or rows of granules on each 
elytron. Ratio of elytral length to width across bases 
about L3;l, Hind wings almost entirely atrophied. 
Longest hind spur equal lo more than Vs. length o( 
metatarsus. Paramcrcs making up about { A of total 
aedeagal length. No sexual dimorphism. Total length 
15-20 mm. 

Distribution and hubmw I he north-east quarter of 
South Australia and adjacent areas of Queensland and 
the Northern Territory. Not recorded from eaves but 
found in rabbit burrows and in ihe open on sand ridges 
at night, sometimes in large numbers. Collected from 
luly to May. 

Tvpe: Killalpanima, S.A., KX) miles east of Lake Eyre 
(H. .1. Hillier), o\ BMNH. 

Specimens examined (119): SOUTH AUSTRALIA. 
Alton Downs (old) H.S., 48 km SW by W of Birdsville. 
Hirdsville, 25 rni S of Clayton R., near bore drain. 
Cooper's Creek. Diamantina R., 25 mi S oi Birdsville. 
Hay R., 24 mi \V of camp 13. Innamineka, Brodie's 
Water Hole. Lake Eyre, Madigan Gulf, Sulphur 
Peninsula. Lake Eyre, Preseott Peninsula. Lake 
Kittakittaooloo, S shore. Lake Palankarinna. Minnie 
Downs, NE corner. Mt. Gason, 41 km SSW of Clifton 
Hills. Mungeranic Sta., water hole. Mungeranie, 20 mi 
S of. Purni Bore, Simpson Desert, 6 km WSW ot\ pit 
traps in sand ridge. Simpson Desert. Warburton R.. 
2 km NE of Kalamurina H.S. Warburton R., New 
kalamunna H.S. NORTHERN TERRITORY. Andado 
H.S., 15 km ENE oi\ J'inke R., McDonnell Ranges. 
Hermarmsburg. Indracowra, 5 mi N of, e\ pit traps in 
rabbit burrows. QULINSI.AND. Arrabury, 20 mi N 



FIGS 15-22 15 Head, Brises nuilarlmricusn. sp. 16. Head, /irises trachynotoides Pascoe. 17 Head, Hnses occutcniuhs n. sp ,18. Head, 
Brises nurvicoliis (Blackburn). 19. Hind femur of <S , ta trachynotoides Pascoe. 20. Middle femur of rf , SmeS trachy notaries Pascoe. 
21. Hind femur of <? , Brises nullarbomus n. sp. 22. Middle femur oi 3 , Arises nultarboncus n. sp. 

of, on road to Planet. Birdsville. Birdsville, 30 km W 
of, Bluff Sla. Diamantina R. Kaliduwarry Sta., Camp 
20. Specimens are located in AMSA, ANIC, QMBA, 

2. Brises caraboides, n. sp. 

(Figs 8, 25) 

Description: Mentiim without pits, with irregular 
surface and narrow anterior membrane. Terminal 
antennal segment subaeuminate. Clypeus in lower 
plane than frons. Eyes somewhat wider than long in 

dorsal view, separated by a distance equal to about 3'/? 
eye widths. Edges o\ prosternal process not raised. 
Pronotum about 1.3 times as wide as long. Pronotal 
surface finely shagreened, with scattered fine 
punctures. Humeri rounded. Elytral surfaces with 
about 15 rows of coarse punctures in straight lines, 
closely spaced between four low, more or less evenly 
spaced rounded longitudinal ridges on each elytron, 
without carinae, with short fine setae. Ratio of elytral 
length to width across bases about 1.9:1. Hind wings 
reduced to about half of elytral length. Longest hind 
spur equal to about Vy of metatarsus. Parameres 


KK . S. AUST MUS. 10(6): 77-*) 

now. tym 

making up about ft of total aedeagal length. Ibtal 
length IS- 1 9 mm. 

Remarks: The species is closely relaicd to acuticornis, 
within whose range it occurs and from which it differs 
mainly in pronotal and elytral shape and puncturation, 
by being (light less, and by having longer legs. The 
narrow humeri indicate that wing atrophy is of long 
standing and no intermediate forms are known to exist 
between caraboidvs and acuticornis. 

Distribution and habitat; The type locality only. 
Collected in an ''alcove at base of eliffs M 

1\pe: Twilight Cove, Eucla Basin, WA , 5.XI.1966, 
J. Lowry, tr. ANIC Puratype: One o\ with same data 
as hulotype, ANIC. 

Brises acuticornis (Pascoe) 

Description: Merit urn without pits, sparsely punctate, 
with posterior V-shaped grooves enclosing raised area. 
Terminal antenna! segment acuminate. Clypcus in lower 
plane rhan irons. Eyes about as wide as long, separated 
by a distance equal to about 4 eye widths. Edges of 
pt05tcrnal process not raised. Pronotum 1.4-1.5 times 
as wide as long. Pronotal surface alutaceous, finely and 
sparsely punctate. Elytral surfaces variable (see under 
subspecies), with short setae. Ratio of elytral length to 
width across bases 1.6-1.7:1, Hind wings Fully 
developed. Longest hind spur equal to about '/» length 
of metatarsus. Paramcrcs making up about % of total 
aedeagal length. No sexual dimorphism, total length 
15-25 mrn. 

I recognize the following two subspecies (taxa 3 
and 4) 

3. Brises acuticornis acuticornis (Pascoe) 
(Tigs I, 3, 26) 

Lphidonius acuticornis Pascoe, [869, p. 15 J. 

Brises acuticornis. Carter, 1914, pp. 45-46; Carter, 
192ft, p. 145; Hamilton-Smith, 1967a, pp. 37-39, 41; 
Hamilton-Smith, 1967b, pp. 115-116; Richards, 1971, 
pp. J 7 -45 passim. 

Description: Elytral surfaces with small punctures 
set in longitudinal rows between three luw ridges on 
each elytron, without granules or spines. 

Distribution and habitat: This subspecies generally 
occurs in the coastal areas of South Australia and the 
Nullarbor Plain. There are numerous records from 
eaves,, from both the light and dark zone, often 
associated with guano and carrion. Probable larvae 
have been collected inside caves and one collector 
(M. Cray) reports that adults and larvae feed on wcta 
(raphidophorid cricket) excreta tn Wcebubbie Cave- 
Collected from July to April. 

In her study of the cavernieulous fauna of the 
Nullarbor Plain. Richards (1971) reports B. acuticornis 
from 25 caves throughout the region, mostly in the dark 

/one up to 4 km from the entrance. Dry, powdery bat 
guano supports large populations of adults and larvae, 
and this species is the only coprophage to occur in both 
bird (swallow) and mammal faeces, including human 
escrcment and fox droppings. It also occurs in rabbit 
burrows on the plain. 

Type: Lectotypc (sex undetermined* from Gawlei, 
S.A., herby designated from two synlypes in the Pascoe 
Collection (HMNH). 

Specimens examined (174): SOU I H AUSTRALIA. 
Ardrossan. Ceduna, Cook, 9 mi E fit Denial Bay, i mi 

5 of. Diprose Cave No. 3. Eucla Basin, cave N 149. 
Fisher, LAY Railway. Fowler's Bay. Gawler. Koonalda 
Cave sinkhole. Koonalda H,S., 10 km SH by E of. 
Koonalda Sta , blowhole entrance. Koonalda, N 33 
sinkhole (amongst dead birds). Koonjbba. 
Kooringabie Sta. Murrawijinee Cave N 7, Nullarbor. 
Murray R. Ooldea. Pt Pierce Mts, Yorke Peninsula. 
Swan Reach, Punyelroo Cave. Thylacine Hole, Eucla 
Basin. Wardang 1. White Wells Cave. Winbirra Cave, 
Eucla Basin. Yorke Peninsula. WESTERN AUST- 
RALIA. AbrakurrieCave N 3, Nullarbor. Caiguna and 
Cocklebiddy, halfway between, ex rabbit burrow. 
Cocklcbiddy Cave, duliue. Dingo Cave N 160, 
Nullarbor Plain, on guano. Eucla. Eucla, 32 mi SE of 
Coongana, Nullarbor. Eyre's Sand Patch. Madura, 

6 mi S of, cave. Madura, 8 mi Cave. Madura, main cave 
N 62. Moonera Tank Cave, Nullarbor. Mullamullang 
Cave, Nullarbor, dark zone, feeding on dead bat. 
Murra-cl-elvyn Cave, Nullarbor, light and dark zones, 
on dry guano. Nullarbor, 50 km W of; 64 km E of; 

7 km WSW of. Nullarbor H.S. Pannikin Plain Cave, 
dark zone. Petrogale Cave, v : mi S of, N 79, E of 
Madura. Swallow Cave, Cocklebiddy, near dead bat. 
Tommy Graham's Cave N 56, Nullarbor Weebubbie 
Cave N 2, Nullarbor. Specimens are located in AMSA, 

4, Brises acuticornis duboulayi (Bates), new status 
(Figs 7, II, 12,27, 28) 

Ephidonius duboulayi Bates, 1872, p. 279. 

Brises duboulayi, Carter, 1914, p. 45; Carter, 1926, 
p. 151. 

Brises granulatus Carter, J92I, p. 316 (new 

Description: Distinguished from a, acuticornis on\y 
by the sculpturing of the elytra, which show at least 
a trace of small granules or spines and an accentuation 
of the ridges. Jn the extreme form, the elytra are 
distinctly spinose and strongly carinate. There is* a 
gradual imergradation between typical a, ucuttcornts 
and a. duboulayi along geographical gradients (see 

Distribution and habitat: This widespread subspecies 
is essentially the inland race of acuticornis, bur ft 
reaches the coast on Eyre Peninsula, where it is also 



in its least granulate form. This form could have been 
ascribed to a. acuticomis, but it is more convenient for 
nornenclaiorial purposes if the latter name is arbitrarily 
restricted to specimens without any trace of granules, 
spines, or carinae. Moderately granulate fornis without 
carinae (the former granulatus. Fig. 27) are to be found 
in eastern South Australia from the Adelaide Hills 
northward and in New South Wales as far cast as the 
foothills of the Great Dividing Range, whilst the most 
strongly carinate and spinose form (a. duhoulayi 
proper, Fig. 28) occurs in northern South Australia 
from Port Augusta northwards to adjacent states and 
westwards through the driest parts of Western Australia 
to the west coast. In parts of South Australia it is 
difficult to assign specimens to one form or the other. 

The Ash ford Cave (N.S.W.) population of this 
subspecies seems to be isolated by many hundreds of 
kilometres from the nearest other records in western 
New South Wales, but the intervening area is too poorly 
collected for us to be sure of this. The few Ashford 
Cave specimens seen are smaller and paler than other 
duboulayi. The other extreme eastern record, Yiddah, 
N.S.W., is based on a single old specimen and needs 

There are several other records of the occurrence of 
this subspecies in caves, and in rabbit burrows (see 
below). Records are from October to July. 

types: Of duboulayi: Champion Bay, W.A., Duboulay, 
<?, BMNH. Of granulates: Broken Hill, N.S.W., R. J. 
Burton, 1/21, 9, MVMA. 

Specimens examined (JOS): SOUTH AUSTRALIA. 
Adelaide. Buckalowie Creek, from small cave a little 
beyond light. Clara St Dora Cave, Cleve, Coober Pedy. 
Evelyn Downs Sta., Oodnadatta. Everard Ranges to 
Warburton Ranges. Franklin I. Kingoonya. Kokatha 
H.S., 17 km SE of, ex pit traps in rabbit burrows. Leigh 
Creek. Minnipa. Mt Finke, 40 km S Malbooma H.S., 
ex rabbit warren. Mt Lofty Ranges. Moralana H.S., 22 
km WNW of. Murray R. Pt Augusta. Pt Lincoln. 
Purple Downs, 160 mi NW r of Pt Augusta. Streaky Bay. 
Stuart Ranges. Tarcoola. Whyalia. Wilgi near Loveday, 
Gilbert's Well. Wilpena Pound, in bat cave in dung. 
Wooltana Cave, 200 ft underground in bat guano. 
NEW SOUTH WALES. Ashford Caves via Inverell. 
Broken Hill. Culpaulin, Darling R. Yiddah. 
TERRITORY. Alice Springs, 15 km S of, bat caves. 


fICi, 23. Distribution ot the species of the genus Bases. I— & hiutrl Carter. 2- B. auvbotdes n. sp. 3 — B. a. aattivontis (Pasvoe). 4 — B 
a. duboulayi (Hales). * — B, kuiherituw u. sp. 6 — B. occidentalis n, sp. 7— tt piirvicoljii (Blackburn). S — B. nul/urhnrivus n, sp. 9 — B. 
tnuhytwioides Pascoc. 250 mm annual isohyet shown. 


RlfC S. ALISI. MUS. tV(fi): 77 90 

./W<r, fflW 

WESTERN AUSTRALIA. Coolgardie. Cue. Onslow. 

Specimens arc located in AMSA, AN1C, SAM A, and 

5, /irises katfwriruie, n. sp 
(Fig. 29) 

Description: Mcntum variable, without .groove but 
with cither large punctures or oblique depression 
posterolateral!}'. Terminal antennal segment acuminate. 
Clypeus in same or higher plane than frons, not 
delimited by a sharp depression, Eyes wider than long, 
the anterior edge strongly oblique, separated by a 
distance equal to about 2 eye widths. Prostet nam feebly 
ridged before coxae, edges of prosternal process 
strongly raised between coxae. Pronotum 1.3-1.5 times 
as wide as long. Pronotal surface irregularly rugose and 
shagreened, vvith numerous large shallow punctures 
and some slightly raised shiny spots on disc. Elytra each 
with three nearly complete carinae and part of a fourth, 
and a sutural row of granules, surface between carinae 
flat, shagreened, with rows of fine but deep punctures 
and sparse short, curved setae. Ratio of elytra! length 
to width across bases 1.8-2.1:1. Hind wings fully 
developed. Longest hind spur equal to about V\ of 
length of metatarsus. Parameres making up about '4 
of total aedeagal length. No sexual dimorphism. Total 
length 15-25 mm. 

Distribution and habitat: Known only from the 
complex of caves near Katherine, NT- It occurs well 
inside the dark zone, and has not been collected from 
outside caves. More collecting in inland northern 
Australia is needed to establish the distribution limits 
of this and any other species in the region, in ordet 
to determine whether katherinac is really as isolated 
as it appears to be. 

Type: Kintore Cave, Kalherine, NT., BS 2134, 
lZv.l974«t& SAM A, I 21 , 155. Para types: Same data 
as holotype. 5, SAMA. Cutta Cutta Cave, NT,, 200 m 
from entrance, S.\ii.I983, W. I). Williams, 2, SAMA 
Cutta Cutta Cave, Katherine, NT. 13. ix. 1973, A. Goede, 
I, SAMA. Three Mile Cave, Katherine, NT. 24.vU%2. 
R, V. Soulhcoll aud NT. Cave Exploration Group, 2, 
SAMA. Kalherine, NX, ! v mile from entrance to 16 
Mile Cave. June 1962, I, ANIC. Katherine, NT., 16 
Mile Cave, 12..v.l%3, W. Penman, 2, WAMA. 

6, times oicidetnalis* n. sp. 
(Figs 9 y 18, 30) 

Description: Mentum with posteto-latcral groove, 
not joining. Terminal antennal segment acuminate. 
Clypens in somewhat lower plane than Irons, but nut 
sharply delimited. Eyes about as wide as lorn?, 
separated by a distance about equal to 4 eye widths. 
Head somewhat elongated behind eyes. Prostenntm 
distinctly earjnate before coxae, edSSS ^( prosternal 
process not raised. Pronotum 1.5-1.6 times as wide as 
long. Pronotal surface shagreened, wilh indistinct 
punctures and very small granules regularly spaced. 

Elytra each with two strong, nearly complete carinae 
and two indistinct ridges externally, intervals between 
them with sparse grannies and punctures in irregular 
rows, without setae. Lateral edges of elytra markedly 
convex. Ratio oT clytral length to width across base- 
1,8-1,9:1, Hind wings fully developed. Larger hind spur 
equal to about W of length of metatarsus. Parameres 
making up about \fi of total acdeagal length. No sexual 
dimorphism, lolal length 19-22 mm. 

Distribution and habitat, Known only from three 
localities in Western Australia situated between 
Meekarhana and ihe coast. One of I he Wdgie Mia 
specimens bears the label "Found in cave". 

type. Weld Range, W.A., 2.iii.l%3, A. Douglas, o*, 
WAMA. Paratypes: Same data as holotype, 4, SAMA, 
WAMA. Mileura H.S. 4 mi W o\\ 8.\ii.l966, W. H. 
Butler, I, WAMA. Wilgie Mia W.A. (27 28 S 114 21 
E), ix. I%1. D. Merilees, 2, WAMA. Ditto, 1%3. A. 
Douglas, 3, WAMA. Ditto, 16.V.1973, M. Thomas, I. 

7 f$ri\e\ pankoilis (Blackburn) 
(Pigs 17, 31) 

Ephidonius parvicoliis Blackburn, 1895, p. 52, 

Brises parvicoliis. Carter, 1926, p. 151. 

Description: Men turn with posrero laleial 
depressions, not meeting. Tetminal antennal segment 
narrowed distally but bluntly rounded. Clypeus in lower 
plane than Irons but not sharply delimited. Eyes large, 
longer than wide, separated by a distance equal to 
about 3 eye widths, Head snrface finely, densely 
granulate and shagreened. Presternum moderately 
ridged before coxae. Pronoium 1.5-1.7 times as wide 
as long, Pronotal surface shagreened with numerous 
very small granules. Elytra each with two sharp but 
fine, nearly complete carinae, and one or two indistinct 
ones laterally. Intervals between I hem shagreened, with 
distinct straight rows of small punctures and minute 
granules, without setae. Lateral edges o\ elytra 
markedly convex. Ratio of clytral length to width across 
bases 1.8-2.0:1. Hind wings fully developed. Longest 
hind spur equal to about ': length of metatarsus. 
Parameres making up about % o\' total aedeagal length. 
No sexual dimorphism. Total length 17-22 mm 

Distribution and habitat: Ihe few available records 
suggest one focus of distribution in the east central part 
v\' South Australia in low-King areas and another in 
the mountains of the north-west corner and adjacent 
ranges o\' neighbouring states. This species has been 
collected from rabbit burrows on several occasions but 
is not known from caves and may be an open-ground 
forager like Main. September to Maieh. 

Type: Lake Callabonna, S.A., A. Ziet/, 9. SAMA. 
I here is \\o specimen by this name in BMNH, The one 
in SAMA is nol marked type but it bears the name 
Ephidonius parvicoliis in Blackburn's w riling and the 



correct locality and collector, and has been designated 
.is lectoiypc. Two oiher specimens with the same data 
are labelled co-types (that is, paratypes) in Blackburn's 
hand, and iherc axe five olhers wilta the Zielz label, 
determined by A. M- Lea as co-types, all designated 
paralectotypcs and all in SAMA. 

Specimens examined (36). SOUTEI AUSTRALIA. 
Agnes Creek Sta. ex pitfalls in rabbit burrows. 
Rradunna H,S,, )5 km W at Evctard Ranges to 
Warburton Ranges. Glenmanyie Bore E of Lake Frome, 
ex rabbit burrows. Lake Callabonna. Lake Mulligan 
(possibly Mulligan Springs near Lake Callabonna). 
Lake Palankarinna (2846 S 138 25 E), rabbit burrows. 
Mora!ana v 22 km NNW of, rabbit burrows.. 
NORTHERN TERRITORY. Petermann Ranges, Hull 
R. 33 km ESE o\' Dorker R. Specimens arc located in 

s. Bases nullarborkus, n. sp. 
(Figs 6, 16, 21, 22, 32) 

Description: Mcntuin with transverse ridges and 
lateral concavities. Terminal antennal segment broadly 
rounded. Clypeus in lower plane than frons, sharply 
set off by a step, Eyes with anterior edges oblique, a 
lillle longer than wide, separated by a distance equal 
to aboul 3 eye widths. Head surface densely granulate. 
Prosternum not ridged Pronotum 1.6-1.7 times as wide 
as long. Pronotal surfaces densely granulate. Elytra 
each with two nearly complete carinae and one short 
humeral carina, rest o\' surface eoaisely retieulo- 
punctalcand granulate, the punctures and granules not 
in tows, without setae. Ratio of elytra! length to width 
across bases 1.8-1.9:1. Hind wings fully developed. 
Longest hind spur equal to about V> length of 
metatarsus'. Paramcres making up about l A of total 
aedeagal length. Sexual dimorphism evident in shape 
of middle and hind femur, the hind margin of which 
is slightly expanded at base in the male. Total length 
13-17 mm. 

Distribution and habitat: Known only from the 
Nullarbor Plain, without habitat data. 

lype: Nullarbor U.S., 7 km WSW of, S.A., 31 28 S 
130 50 \i., D. C. F. Rent/ and M. J. D White, 
Slop 25. 9 , ANIC. Paratypes: Same data as holotype* 
», ANIC. Nullarbor H.S , S.A., 1011960, P Aitken, 
I. SAMA. Kooringibbie (Well), S.A.. 3, SAMA. 744 
Mile Camp, EastAVest Kali way, W.A., F Mack, J, 
SAMA. Fisher, Last-Wcsi Railway, S.A., Troughton and 
W light, 1, AMSA. Reid, 40 mi N of, W.A„ 4 ii 1968, 
S9/3 A. M. Richards, L ANIC. 

9. Brises (rachynowides Pascoe 
(Fig.s 5. 10, 13. 14. 15. 19, 20, 33) 

Brise* trachynotoides Paseoe, 1869, p. 146, pi. xi, 
fig. 5; Carter, 1914, p. 46; Carter, 1926, p. 151. 

Description: Mcnturn with indistinct posterolateral 
depressions Terminal antennal segment broadly 
rounded. Clypeus in same plane as frons. Eyes with 
anterior edges nearly transverse, wider than long, 
separated by a distance equal to about 2'z eye widths. 
Head surface vermieulate Prosternum slightly ridged 
before coxae. Pronotum 1,6-1.9 rimes as wide as long. 
Pronotal surface vermicuiate-granulale. Elytra each 
with two sharp, nearly complete carinae, no humeral 
carina, rest of .surface reticulo-punctatc and granulate, 
the punctures and granules not in rows, without setae. 
Rriiio of elyiral length to width across bases 1.6-1.8:1. 
Hind wings fully developed. Longest hind spur equal 
to about 2 A of length of metatarsus. Paramcres making 
up -about '/* of total aedeagal length. Sexual 
dimorphism evident in shape of legs, the males having 
the hind margin of the middle and hind femora 
angularly expanded and all tibiae distinctly arcuate 
(straight in the female). Total length 10-17 rnm. 

Remarks; There is considerable variability both in 
size and shape in this species, the larger individuals 
having more explanate and sinuate pronotal margins. 

Distribution and habitat: A widespread species 
occurring in Ihc northern half of South Australia, 
south-western Queensland, the south of the Northern 
Territory and all of Western Australia except the South 
West. It is not frequently encountered, however, Found 
in rabbit burrows, and there is one scries from a cave 
on Barrow Island, W,A. August to May, 

Type: Champion Bay, W.A., ?, BMNH. 

Specimens examined (52): SOUTH AUSTRALIA. 
Agnes Creek, ex pitfalls in rabbii burrows. Ediacaia. 
Emu, 300 mi NW of Woomcra. Everard Ranges to 
Warburton Ranges. Mt Finke, 40 km S of Malbooma 
H.S,, e\ rabbit warren, Purni Bore, Simpson Desert, 
6 km WSW of, pit traps on sand ridge. Stuart Creek 
H.S* 10 km E of. Wynbring. QUEENSl AND. 
Hermannsburg. idracowra H.S,, 5 mi N of> es pil traps 
in rabbit burrows. MacDonnell Ranges. Petermann 
Ranges. WESTERN AUSTRALIA. Barrow L cave N 
of Flacourt Bay. Forrest River District. Gill Pinnaele, 
Mural Crescent. King Sound. Kookynic. Mt Unden, 
7,5 km NNW of, Specimens are located in ANIC, 


I wish to thank the curators of the collections 
consulted for arranging loans of specimens, in 
particular Mr M. J. D Brendell (British Museum), Mi 
G. Holloway (Australian Museum), Dr T. F. Houston 
(Western Australian Museum), Dr J. F. Lawrence 
(Australian National Insect Collection). Dr G. B. 
Monteith (Queensland Museum ) % and Dr A. Neboiss 
(Museum ol Victoria, Division of Natural History and 


REC. S. AUST. MUS. 19(6): 77-90 

June, 1986 

FIGS. 24-27. 24. Brises b/airi Carter,d\ 25. Brises caraboides n. sp., 6 . 26. Brises a. acuticornis (Pascoe), <S . 27. Brises a. duboulayi 

(Bates), <$ , granulate (eastern) form. Photos: J. Forrest. 



FIGS. 28-31. 28. Brises a. duboulayi (Bates), <$ , earinate (western) form. 2S). Brises kutherinae n. sp., t? 

d\ 31. Brises parvicollis (Blackburn), <$ . Photos: .1. Forrest, 

30. Brises occidentalis n. sp., 


RFC- S. AUST. MUS. 19(6): 77-90 

Jit tie. IP86 


CARTER, H. J. 1914. Revision of ihc subfamily Tenebnoninae, family 

"Tenebrionidac. Proc. Linn Sac. N.S.W. 39; 44-K6, 
CARTER, H. J. 1926. A check list of the Australian Tenebrionidac. 

Aust. Zooi 4: 117-163. 
DOYEN, J. T, and I AWRENCE, J, F. 1979, Relationships and 

higher classification of some Tcncbrionidae and Zopheridae 

(Coleoptera). Svst. Ent. 4: 333-377. 
DOYEN, J. T. and TSCH1NKEL. W. R. 1982. Phonetic and eladistic 

relationships among tenebrionid beetles (Coleoptera). Svst. Ent. 

7; 127-183. 
HAMILTON-SMITH, E. 1967a. Fauna of (he Nullarbor Caves, In 

"Caves of the Nullarbor" (J. R. Dunkley and T. M. U Wigley, 

cds.), pp. 35-42. Speleological Research Council Ltd., Univ. of 

Sydney. 61 pp. 
HAMILTON-SMITH. E. 1967b. The Arlhropoda of Australian caves. 

J. Aust. ent. Soc. 6: 103-118. 
KASZAB, 7. 1982. Die lenebrioniden Neukaledoniens und der 

Loyaute-Inseln (Coteoptera). Folia ent. Hungurieu 43: 1-294. 
MAI THEWS, E. G. 1985. Foraging activity of some tenebrionid 

beetles in a South Australian malice area. In "Soil and Litter 

Invertebrates of Australian Mediterranean-type Ecosystems" (P. 

Greenslade and J. D. Majer, eds.). H'A Inst. Tech., Sch, Biol. 

Hull. 12: 63-64. 

MEDVFDFV, G. S. 1968, "Zhuki chcrnotclki (Tcncbrionidae). 
Podsemeistvo Opatrinae^ Fauna SSSR Vol. 19 t pari 2. 

MEDVEDEV, G. S. 1977. Taksonomicheskoe /nachenie 
antennal'nykh sensill zhukov-chernotelok (Coleoptera, Tcnc- 
brionidae). 7?. Vsesoyttzn. ent, Obshch. AN SSSR 58: 61 86. 

PASCOE, F. P. 1869. Descriptions of new genera and species of 
Tcncbrionidae from Australia and Tasmania. Ann, Man, Nut. 
Hist. 3: 29-45, 132-53, 277-96, 344-351. 

RICHARDS, A. M. 1971. An ecological study of the cavcrnicolous 
fauna of the Nullarbor Plain, Southern Australia. J. Zooi. Land. 
164: 1-60. 

TSCHINKEL, W. R. t and IX)YEN, .1. T. 1980. Comparative anatomy 
of the defensive glands, ovipositors and female genital tubes 
o\' tenebrionid beetles (Coleoptera). Int. J. Insect Morphoi 
Embryoi 9: 321-368. 

WATT, J. C. 1968: Specific synonymy in Mimopeus Pascoe {Ctlihe 
auctonim), and the nomenclaloral status of some related genera 
(Coleoptera, Tenebrionidae). N.Z. Ent. 4: 35-39. 

WATT, J: C. 1974. A revised subfamily classification of Tenebrionidac 
(Coleoptera)- N.Z. J. Zooi fc 381-452. 

WALT, J. C. 1979. Abbreviations for cntomotouical collections, N.Z. 
J. Zooi 6: 516-520. 

FIGS. 32-33. 32. Htises mttlarhoricus n. sp., 9. 33. Buses truchynoto'ules Pascoe, cf . Photos: J. Forrest. 


by Penelope Greenslade 


The fauna now consists of 8 species and 2 doubtful records. Friesea tilbrooki, Cryptopygus caecus, 
Cryptopygus tricuspis, Isotoma (Sorensia) punctata and Isotoma sp. indet. Are new records for the 
island, and the presence of Tullbergia bisetosa and Cryptopygus antarcticus antarcticus is 
confirmed. Tullbergia templei was described from Heard I. As was Friesea viennei which is here 
synonymised with Friesea tilbrooki, and Cryptopygus quadrioculatus is synonymised with C. 
tricuspis after examination of types. Records of Isotoma (Sorensia) subflava and Parisotoma 
octooculata from Heard I. Are no longer valid since they depend on incorrect identifications and 
Tullbergia antarctica and Cryptopygus antarcticus reagens are considered doubtful records. The 
fauna consists mainly of widely distributed species and there is no sign of endemism. 




Honorary Research Associate, South Australian Museum, North Terrace, Adelaide, South Australia 5000 

{Manuscript accepted 30 April 1985) 


GRLLNSI \l»-. I'KNH OPE ls)H6, Additions tO the collembotan 
Imuih Of fattti fsh.nU. AYt. S. Aust MUX. I9(7|; 01-06, 

The fauna now consists of 8 species and 2 doubtful 
records. Friesea tilbrooki, Cryptopygus caecus, 
Cryptopygus tricuspid Isatoma (Sorensia) punctata 
and Isatoma sp. indet. are new records for the island, 
and the presence of Tultbergia bisetosa and 
Crvptopygus antaredcus antarcticus is confirmed. 
Tullbergia templet was described from Heard I. as was 
I nesea viennei which is here synonymised with Iriesea 
til brook i, and Cryptupygus quadrioculatus is 
synonymised with C. tricuspis after examination of 
types. Records of Isotoma (Sorensia) sub/lava and 
Parisotoma octoocula/a from Heard I. are no longer 
valid since ihey depend on incorrect identifications and 
Tultbergia antarciica and Cryptopygus antarciicus 
reagent are considered doubtful records. The fauna 
consists mainly of widely distributed species and there 
is no sign of endemism. 


Heard I. is a small (43 kmXl9 km) ice-capped 
Antarctic island lying in (he South Indian Ocean at 
53°06'S ( 77°30 E aboul 440 km SE of the Kerguelen 
Is and 4100 km WSW of Perth. Parts of the island are 
permanently glaciated and it is dominated by a 
quiescent volcano, Big Ben, which rises to 2745 m (Law 
and Burstall 1953; Home 1984). Other areas near the 
coast and on headlands arc free of ice and snow for 
sonic months of the year and support vegetation. Seven 
vascular plants have been recorded. Several collections 
of Collcmbola have been made from the island 
(Table 1). Wise (1970b) dealt with the fauna in detail 
and listed seven species while the most recent review 
is that Of Ueharveng (1981) who did not examine 
previous collections and gave three definite records and 
five doubtful identifications. A new collection of 
Collcmbola from Heard I. was found to contain two 
species not recorded before and this prompted an 
examination of material collected previously and a 
review of the faunal list. Identifications were made 
using Dcharvcng's (1981) keys, descriptions, figures and 
specimens identified by him unless indicated otherwise. 
Subspecies have heeu retained although this should not 
be taken as support by the author for the concept 










Vanhoften 1V0S 
1901 3 iimlcilein 1900 Ikrljn? 

Deutsche Slid 
Polar Hspcdi- 


BAN/AKL 1929 Womerstev 1937 SAMA 

(V. H, Juhnvhnil 
AKAKE 19510 Brown |%4 L'SPHTM 

(K, G. Brown) 
V, Temple 1965 Wise 1970b ANIC Ihololypcs) 

S. Trcmoni 19X3 Trojuout 1983 SAMA 

this work 

The following abbreviations are used: AMNZ, 
Auckland Institute and Museum, New Zealand; ANIC, 
Australian National Insect Collection, Canberra; 
BAN/ARE, British, Australian and New Zealand 
Antarctic Research Expedition; BPBM, Bern ice P. 
Bishop Museum, Honolulu, Hawaii; LDCoIl, Louis 
Dcharveng Collection, University Paul Sabatier, 
Toulouse, France; NMNZ, National Museum of New 
Zealand, Wellington; SAMA, South Australian 
Museum, Adelaide; USPHTM, Department of Public 
Health and Tropical Medicine, Sydney University, 



1 Thora* I carrying setae, not reduced, abdomen V and 

V t separate 2 

Thorax J without setae, reduced, abdomen V and VI 
fused , - . ... 5 

2 White, ocelli absent, two spines posterodorsally on 

abdomen V|, grinding mandibular plate and pscudo- 

celli present, furca absent 3 

Grey, ocelli 3 + 3, 7-11 spines posterodorsally on 
abdomen VI, no grinding mandibular plate or pseudo- 
celli, furca present Friesea tilbrooki Wise 

3 Empodial appendage present with seta at least half 

as long as claw ....... Tultbergia bisetosa Burner 

Lmpodial appendage absent or rudimentary, lacking 
seta 4 

4 Postamenna! organ with 50 tubercles or less, small 

species, less than 1.5 nun long 

Tullbergia templei Wise 
Postantennal organ with 80 tubercles or more, large 

species, more than 2 mm long . . . , 

TuttbetRtu unturcticu Lubbock 

5 White, ocelli and pigmented eye patch absent .... 6 
Greyish or black, ocelli and pigmented eye patch present 



REC. S. AUST. MUS. 19(7): 91-96 

Ju/ie, t9H6 

Dens aboul 3 times Kjngei than manubrium, with more 
lhan 15 anterior setae and more than 6 posterior setae, 
mucro with three teeth lsotomu sp, intlet. 

Dens aboul the same length as manubrium, with 9-13 
anterior and 4-5 posterior setae, muero wilti 5 teeth 
Cryp/opyuus meats Wahlgren 

Ocelli 2 I 2 or less, no clavate tcnent hairs, muero with 
3 teeth - 8 

Ocelli (H 6, 2 da\ule lenent hairs present on all legs, 
mucro with 2 teeth 9 

While with sparse black speckles, ocelli 1 t- 1, dens nearly 
.V- longer lhan manubrium with about 8 posterior 
and more than 50 anterior setae, body with long ciliated 
macroehaetae distally abundant about 4 * longer than 
ordinary setae on abd V/VI, distal inner anterior 
margin of the manubrium with 1 » I spines with swollen 
hases .... Isotoma (Sarensia) punctata Wahlgren 

Grey, ocelli 2 + 2. dens with 5-6 posterior setae and 13-15 
anterior setae, dorsal macroehaetae smooth and aboul 
2 x as long as ordinary setae on abdomen V/Vl, distal 
inner anterior margin of the manubrium without spines 
Cryptopy^us tricuspid Enclerlein 

Dens shorter than manubrium with 5 anterior setae and 
four posterior setae, mucro with two teeth, macro- 
ehaetae smooth and only about a third as long again as 

ordinary setae on abdomen V/VI 

Cry ptopy gits antarctictts Willem 

5 (rarely 4) poslcrior subcoxal setae on fureal segnieni 
and 15-18 anterior sub-eoxal setae, 11-18 posterior 

manubnal *elae 

....... CryptOpygtts uitturcficus rcaxens Fndcrlcin 

4 (rarely 5) posterior su banal setae on luteal segment 
and 8-14 amerior sub-coxal seise, 9 + 9 (range 8-10) 
posterior manubnal selae , 


I friesea tilbrooki Wise 1970 

- Friesea viennei Deharveng 1981 syn. nov. 

(Tigs I, 2) 

Type Locality: South Georgia. 

Material Examined: Holotype: South Georgia, 
Busen Peninsula, Enten Bay, under rocks on beach, 
7.xi.l963, H. D. Clagg, SG 28A, KWSG 216c (BPBM); 
Heard 1., Mt Aubert de la Rue, grassy lawn, 26.ii. 1941. 
Dr Vienne, KWHI 39, (Holotype of Friesea viennei) 
?LDCol|.; Macquarie L, various localities, 32 exs. 

Wise (1970a) described F. tilbrooki from three 
specimens apparently with 10 anal spines collected on 
South Georgia. In 1981 Deharveng described E viennei 
from Heard I. from a single specimen with seven anal 
spines. He mentioned that F, viennei was close lo E 
tilbrooki differing only in the number of anal spines 
and in having at least four sensory setae (soies s) on 
antenna IV compared to E tilbrooki's three. Re- 
examination of the type of F. tilbrooki shows that it 
possesses five sensory setae (Fig. 2) \x] this position and 
that in all other respects apart from the anal spines it 
is identical to Dcharvcng's description of E viennei. 
The two paratypes ol E tilbrooki have recently been 
examined and possess 7 and 8 spines respectively on 
abdomen VI (K. Wise pcrs. eomm.). Friesea viennei 

was described with a small tooth to the claw and 
although E tilbrooki lacks this tooth according to Wise, 
a small looth was observed by the present author on 
claws I and II of I he / tilbrooki holotype. 



a a 

. 6 



33 optteirnsnt 







10 apeqimona Unm c 

» 1 

8 9 to 

Numoor Of apin»a 

FlO. 1. Number of spines on .abdomen VI found on f'riesca tithraokt 
Wise specimens from a number of localities on Macquarie 1. 

Morphological variation: Examination of a long 
series of specimens of E tilbrooki from a single locality 
on Macquarie I. showed that the number of anal spines 
ranged from 7-11 in ad litis (Fig. 1) and that this 
variation had a similar distribution wiihin a single 
population to that found for all specimens examined. 
A third of the specimens were asymmetric and seven 
was the most common number of spines with aun ( m : 
and p„ always spinose. The holotype of E tilbrooki 
from South Georgia had m,. and a spinose also (Wise 
1970a, Fig. 2E); one of the paratypes was asymmetric. 
A single specimen from Macquarie I. had II spines 
(Fig. 2E). No correlation was found between age, Stee 
or sex of specimens and the number of anal spines. 
Bifurcate and double spines inserted in a single position 
were seen occasionally. 

The sensory setae on antennae JV varied in si/c in 
adults. Occasionally individuals with 4 or 6 well 
developed sensory setae were found. The tooth on the 
claw was small and hard to distinguish. It appeared to 
be absent in some specimens and on some legs, 

Other variation found in the Macquarie I. 
specimens was in the tcnent hairs from fairly short to 
long and slightly clavate or bent at the lip. The length 
of the abdominal macroehaetae varied also and when 
longest they were slightly serrated and bent at the tip. 

Deharveng mentions that F viennei has a tendency 
to plurichaetocity and was asymmetric in its setal 
arrangement particularly on ihorax II and abdomen 
IV. He compared its plurichaetocity to Friesea fara 
which was studied by Grow and Christiansen (1974). 



FIG. 2. Frieseu (i/brooki Holotype, A. dorsal view of antenna IV (sensory setae stippled), B. mandible, C. maxilla, D. genital opening 
OJ'S specimen from Maequariel., F. dorsal view of abdomen VI, (cuticle only partly drawn). F. genital opening of d" . Scale line = 0.1 mm. 


REC. S. AUST. M(JS. 19(7); 91-% 

June, 19X6 

These ambers noted that another species, Friesea 
grandis, had a Remarkable amount of setal variation"* 
and that Specimens were rarely symructrical. Scries of 
specimens of F. lilbrooki from Macquarie I. also 
showed much variation in setal number and 


111 -Mil) IS| ANU \N1) MA(.Qi:ARIF IS! ANO 

I Macquarie 1. Spec i men »* 
Nurnbft of specimen^ - 17 


A dorsal leneiH hjir III I til- claw III - 1,062 mean * - .255 

95^0 CI. 1.17, .45 ran*: .75-1.45 

^ = -1153 
B Int claw. III: diam oc A 3.2 mean i -.94X1 

95°-'o C.I- I*, 2.8 aum: 2J-5.7 

7 .14N4 

Canal *pincp a . Int. claw 1IU.81I mean v =.19 

WVo C I. 89, 73 range 56-L4 

v -.2950 

II CompyriMin GJ Ratios 

A B C 

Holoivpe /. vierwei (Heard I ) 

(according to Deharven^ (19810 1-25 3 7 V (0.7-0.9) 

Hololypc F.~tilhrooki (S. Georgia) ,146 2.5 Rfcrt 

Maapiarie I specimens ' 1.06 3.2 0.811 

Measurements of the dorsal tenent hair and internal 
claw length of leg III, diameter of ocellus A and anal 
spine p„ were made and ratios of their lengths 
compared with those given by Deharveng (1981) for l\ 
viermei (Table 2). Dcharveng's value of 1.25 for ratio 
A (dorsal tenent hair: internal length of claw lit), is 
higher than the mean found here but within the range 
of these observations. Ratio B (internal length of claw 
III: diameter of ocellus A) he gives as 3 and ratio C 
(anal spine: internal length of claw Ml) as 7-9. The 
latter figure is certainly a misprint for 0.7-0.9 which 
falls well within limits found here. This is good 
agreement considering the variation in size of different 
spines on a single animal and that Deharveng does not 
state which spine he measured. 

In view of the measured variation within the 
Macquarie I. populations and known variation in other 
Friesea species (Grow and Christiansen 1974) J consider 
the specimens from Macquarie I., Heard I. and South 
Georgia to be conspccific. 

No other southern hemisphere species of Friesea has 
the same combination of eye number (34 3), anal 
spines (7-1 1 ) and well developed furca. hriesea 
multispinosa Denis 1947 is nearest, described as having 
14-15 spines (13 in Denis 1 figure), 3 + 3 ocelli and with 
a similar furca. Denis (1947) based his description on 
a single specimen from Kerguelen Island found in a 
similar littoral habitat in which F. iilhrooki is found. 
In view of the considerable variation in the number of 
abdominal spines of F tilbmoki and its similarity to 
F multispinosa in habitat, ocelli number and furca, it 
seems possible that it is a synonym of F, multispinosa. 
However until the holotype of F multispinosa is 
examined no change of status of these two species is 

Some additions to the description o\ F, iilhrooki are 
given below, although details given by Deharveng and 
by Wise, apart from those already corrected above, still 

Antenna IV: 5 well developed sensory setae (soics s). 
small organite and accessory sensory seiae present, 
single apical bulb with slighl suggestion of division. 

Mouthparts: mandible with 7 teeth; maxilla with 
dentate lamella carrying about 12 teeth. 

Genital Aperture: 9 with up to 30 short setae and 
one pair on anterior lip of genital opening; cr with 4 + 4 
broad round sensillac internally and about 50 short 

Chaetotuxy: abdomen VI with tendency towards 
plurichactocity and asymmetry, a t m t m ? p„ (practically) 
always spinose, m ( , usually absent 

Comments on the biology of this species and details 
of Macquarie J. collections will be given in a later paper 
(Greenslade and Wise unpubl. results). 

Distribution: Heard I, Macquarie L, South Georgia. 


2. Tidlbergia antarctica Lubbock 1876 

Type Locality: Kerguelen J. 

First recorded for Heard I. by Vanhotfen (1908) and 
linderlein (1909). This record was noted by Salmon 
(1949), Brown (1964) and Wise (1970b) but it has not 
been recollected. 

The specimens have not been found. This is a 
doubtful identification according to Deharveng (1981). 

Distribution: Antarctica, Kerguelen I. 

3. Tullber%ia bisetosa Burner 1903 

Type Locality: Kerguelen Is. 

First recorded for Heard 1. by Womerslcy (1937). This 
record was noted by Denis (1947), Salmon (1949), and 
Deharveng (1981). The species was recollected by Brown 
and by 'Fremont. 

Material Examined: Heard I., Skua Beach, detritus 
under rocks, in Azorella and in rock crevices, 18 ii. 1983, 
S, Tremont, 26 exs (SAMA); Atlas Cove, beneath Poa 
near elephant seal wallows, 20.ii.i983, S. Tremont, 50 
exs (SAMA); Atlas Cove, beneath rocks, in Azorella 
near elephant seal wallows, 20.ii.1983> S. Tremont, 31 
exs (SAMA), Heard 1, BANZARE coll. 356, del. H. 
Womersley, 3 exs (SAMA); 31 slides (about 100 ex-,) 
K. G. Brown, no further data, (USPHTM). 

Distribution: Kerguelen Is, Macquarie J. ( Heard I. 

4. Tultbergia templet Wise 1970 

Type Locality: Heard I. 

Nor recollected but record noted by Deharvent: 

Material Examined: Holotype: Heard I., & Barrier, 
Cairn 5, 457 m, #Ji.l965, R Temple, <ANIC). 



Distribution: at prcsenl only known from Heard I. 
and Vlaequarie I. (Greeushide& Wise unpubl. results). 
Dchat veng (1981 ) recorded a species from the Kerguelen 
Is which he identified tentatively as Tultbergiu ef. 


5. Cryptopygus antarcticus antarcticus Willem 1301 

Type Locality: Antarctica. 

F 1 1 s I record cd lb c H ea rd 1 . as Crypfopygus 
antarcticus by Wornersley (1937) and recollected by 
Brown, Temple and Tremont. 

Material Examined: Heard I., Atlas Cove, beneath 
Poa near elephant seal wallows. 20. ii. 1983, S, Tremont, 
2 exs, (SAMA); beneath Azorella on hillside, 1,12.1929. 
BANZARE coll. 349, del. H. Wornersley, 3 exs 
(SAMA); 12 slides (about 20 speeimens) K. G. Brown, 
no furl her daia (USPHTM). 

Distribution: Widespread in Antarctica and 
Subantarclic islands, ?New Zealand, ?Australia. 

6. Cryptopygus antarcticus rcagens Enderlein 1909 

type Locality: I. 

First recorded for Heard I. by Vanhoffen (1908) and 
Enderlein (1909) as Cryptopygus reagens Enderlein. 
Not recollected although the record was noted by 
Brown (1964) and Wise (1970b). Dcharveng (1981) 
considers this to be a probable misidenlifieaiion for 
C. a. antarcticus Willem. The specimens have not been 

Distribution: Cro/et I .. Possession J ., lie des 

7. Cryptopygtis caectts Wahlgren 1906 

type Locality: South Georgia. 

First collected from Heard I. by Tremont. 

Material Examined: Heard I., Skua Beach, detritus 
under rocks near Azorella and in rock crevices, 
I8.ii.1983, S. Tremont, 38 ess (SAMA); Atlas Cove, 
beneath Poa grass near elephant seal wallows, 
20.ii.l983, S. Tremont. 5 exs (SAMA). 

Distribution: widespread in S. America, Australia* 
New Zealand, Subantarclic islands and Antarctica. 

8. Cryptopygtis tricuspis Enderlein 1909 
- Purufolsomui quudrioculata Wise 1970 
-Cryptopygus quadriocuiatus Wise 1974 nee 
(Kapoport 1963), nee (Marlynova 1967)* 
* The two different species described by Rapoport and 
Manynova and named fwfomina quadrtocuiata were 
automatically transferred to the genus Cryptopygus 
when Isotomitla was synonymised with Cryptopygus 
by Massoud and Rapoport (1968). 

Type Locality: Kerguelen Is. 

First collected from Heard I, by Tremont. 

Material Examined: Heard ]., Atlas Co\c, benealb 
Pou grass near elephant seal wallows, 20.ii.1983, S. 
Fremont, 2 exs (SAMA); Atlas Cove, beneath rocks 
neat Azoreliu, 20.ii.1983, S. Tremont, 1 ex (SAMA). 

Deharveng (1981) notes that C tricuspis from Kerguelen 
I. and Wise's C quadriocuiatus from South Georgia 
are very close except that C. quadriocuiatus has only 
9 'ventral' (anterior) setae on the dens and that 
according to Wise's text the species seemed not to have 
differentiated macrochaetae on the thorax or abdomen 
except at the extreme posterior end of the body. 
Examination o\' the holotype of Parafolsomia 
quadriocutata showed that both denies carried 13 
anterior and 6 posterior setae. Macruchaetae were 
present as follows; on thorax II t, thorax 111 1, 
abdomen 1 3, II 3, III Z^ropty sockets and IV 
3 4 empty sockets. This pattern conforms with lhat 
given by Deharveng. The macrochaetae seem to be 
easily detached. A paratype carried 12 + 5 and 13 + 5 
setae on the denies. Other details of chaetotaxy were 
as follows: 

Numbers of setae 

C tricuspis P. P. 

del quadrionttuta quadnvcuhua 

Deharveng Holotype Paratype 





P ureal subcoxa 



14, 17 

17, 21 




6, 6 

In all other respects the holotype and a paratype of 
Parafol-sotnia quudrioculata agree with the 
description given by Deharveng (1981) for C tricuspis, 
and 1 therefore consider them conspccific. 

Distribution: Kerguelen Is., Crozct ]., Marion I., 
South Georgia, Heard I 

9. Isotoma (Sorensia) punctata Wahlgren 
Type Locality: 1 "terra del Fuego. 

Material Examined: Heard l. f "from damp 
situation". Lxii.1929, BANZARE Coll 356, 1 example, 
labelled Isotoma octo-oculata Willem, det. H. 
Wornersley; Heard I., Poly Gully, from dove prion's 
(Pachyptila desalata) nesting material, 8.ii.l965, P. 
lempie KWH121, about 19 exs (BPBM); 2 slides (3 exs) 
K. G. Brown, (labelled Parisotoma octo-oculata), no 
further data, (USPHTM). 

This species was recorded by Wornersley (1937) and 
Brown (1964) as isotoma octooculata Willem 1901. 
Wise (1970b) recorded Temple's specimens as Sorensia 
subflava Salmon (now Isotoma (Sorensia) sub/lava 
according to Deharveng (1981)). Material has been 
compared with the holotype of /. (S) subflava and 
found to differ from it in a number of characters of 


REC S, AUST. MUS. 19(7): 91-96 

June. 1986 

specific importance; i.e. sensory setae on abdomen V 
and VI, and chactotaxy of the manubrium. Specimens 
are similar to /, (Sj punctata in These characters and 
are identified as this species. A revision of the genus 
is in preparation. 

Distribution: Tierra del Fuego, Crozet I., Possession 
[., Argentina, Heard I. (ail other records of this species 
require verification). 

10. Isotoma (Parisotoma) octoocutata Willem 190! 

Tvpe Locality; Antarctica. 

This species was determined by Womcrsley (1937) 
and Brown (1964). Both Womcrslcy's and Brown's 
specimens are now found to belong to /. {$) punctata 
(sec above). Wise (1970b) noted the species record but 
Deharveng (1981) considers the record of this species 
doubtful. There is clearly now no evidence to suggest 
it is present on Heard I 

Distribution: Antarctica only (all records of this 
species from Subantarctic islands, such as South 
Georgia (in Wise 1970a), require re-examination). 

II. Jsotoma sp. indet. 

Material Examined; Heard I., Poly Gully, Winston 
Lagonn, P. Temple, 2.ii.l965, KWHI2, 3 exs (BPBM). 

These three specimens were identified by Wise 
(1970b) as Sorensia subftava. They are in poor 
condition and it is not possible to determine them 
furl her than to genus. They are unlikely to be /. (P) 
octooculata since the postantennal organ and 
ciiaetotaxy of the manubrium differ from those of that 
species. Similarly the structure and chactotaxy of the 
manubrium differ from /. fS) punctata. 


Of the eight species definitely recorded, six occur 
widely on Subantarctic islands and Lhe seventh, T. 

temptei occurs on Macquane I as well as Heard I. 
(Grcenslade & Wise unpubl. results). One species is of 
unknown distribution. No island endemism was detected. 
Deharveng (1981) noted some island endemism for the 
Kerguelen Is, but the fauna ol these islands is richer 
in species, and the islands are larger in area and further 
north. However the fauna of Macquarie I. with 22 
species also shows little island endemism (Grcenslade 
and Wise unpubl. results) and it is al about the same 
latitude and of the same si?c as the Kergueleiis and 
near the rich source area of the Auckland and 
Campbell Islands. 

A large amount of morphological variation, 
particularly in chactotaxy, was found in these Sub- 
antarctic species. Other invcrtcbralcs from polar regions 
have been found to exhibit a high degree of variability 
(Downes 1965). It has been suggested by Grcenslade 
(1983) dial this is because these faunas arc subject 
to A selection where relaxation of consirainls 
on conservative morphology occurs. Whether the 
variation documented here is due ro a genuine 
divergence of lineages on different islands, or 
to relaxation of selection pressures or even to the 
founder effect can probably only be determined by 
breeding, eytological or biochemical techniques. 


1 would particularly like to thank K. A. J. Wise of 
i he Auckland Institute and Museum for much help of 
various kinds during the preparation o\ this paper and 
for critically reviewing the manuscript and also for the 
loan of specimens. Thanks are also due to Dr R. G. 
Ordish, National Museum, Wellington, Professor J. 
Brian of the University of Sydney's Department of 
Public Health and Tropical Medicine, L Deharveng, 
University Paul Sabatier, Toulouse, and the Bcrnice P, 
Bishop Museum for the loan of specimens and to the 
British Trans-Antarctic Fund for financial support. 


BROWN. K C. \%4. The insects ol Heard Island. ANARL Rep. 

Ser. U Vol 1, /ooloev. Pub! No. 7J- t-J9. 
DEHARVENG, L. 1981. Collemboles des ties subuntarciinues de 

I'Ocean Indien. Com tie national ftan<;ais fifes ivcfu'trhx.'^ 

aniao-fiques, Bioloiih i7e5 Sofa Mo. 48: 33*108 
DENIS, t t K. 1947. Collemboles. Croisiere du Bougainville aux Zips 

australcs Mem. da. Mus. Hist. S'ai, Pans Jnjr.J 20: 

DOWNKS, .). A. 1965. Adaptions of insects in lhe Arctic. Am 

AW. em 111: 257-274. 
FNDERl EIN, O. 1909. I0j- Inseklcn Ota AnlarM.scheu Gchieics. 

Deutsche Sudpolar Exped. I90M903. 10: 36^528, Berlin. 
GKLINSI \OL, P, | ML 1983. Adversity selection and die habitat 

templet Am Nul 122: J52-365. 
GROW, A- &., and CH« ISTIANS1N. K, 1974. Ch;iemifi\.v in 

ncarctic Pncseti (Collembola Ncaniirinae) with notes on 

taxonomic u\t J o! ehaetotaw. AVv. fu-ot, Biol. Sol. U: 377-396. 
(K)RNb, K A. J9N4. Terrestrial invertebrates from VlaeHonald Island 

.'iml HcmO U.i.kI, S.ih-AnUKCUv- /. Attst. era Sffc 23; 3« 

LAW, P. (J., and BURSTAI L. T 1953. Ilcaid Island. ANARl 

Interim Rep. No 7_ I'ulr No. 12: 1*32. 
MASSOUD, /.., and RAPOFORT. K H. |«6S C t»ll wholes 

isotomide-, d'-Xmerique du Sud et de r\ji!div»k|ik. Mot /' \nu r. 

(tl0f t |V 307-33r 
5AJ MON, J. T. 1949. New sub -antarctic C nllcrnbola. C<//>r /f.v/w/. 

Sir, But/. 4; 1-56. 
I Rl MON T, S, 19K3, ( "ollcmbola. In Heard Island i xpcdirton 19M3 

Scientific Report fkl, R, Vining. p. 31. 

VAMHOI U:N. U. 190K. Tim- und PfUnzen der HtarcMnscl. 

Deutsche Sudpolacl-xped I9OI-I903, 2: 2o5-2"l, Berlin 

YATSlf. K A. X 1970a. Collembola of South Georgia t\/t.iiit lusters 
Monograph 1M IH3-20H. 

WISt, k. A. I. 1970b. Collembola of Heard island, fthtfu tot. 7s 

Monomwh 23: 209-21?. 
WUMIRSLKV, H. 1937 Collembola (Spitn^tnilsj. tiA : \/.\RLK^\ 

Ser. U, Vo!.4: l-7_ 



byW. ZeidlerandK. L. Gowlett 


The South Australian Museum collection of chiton types is the largest in the Southern hemisphere. 
It contains primary type material, and some secondary types, of 123 species, subspecies or varieties. 
A further 15 species are represented only by secondary types. The type status of at least 11 of the 
primary types is considered very doubtful or, in the case of neotypes, invalid. Listed in an appendix 
are an additional 21 specise represented by types of dubious status from the Dupuis collection. 
Species are listed alphabetically according to the original name of the genus or species. 




South Australian Museum, North Terrace, Adelaide, South Australia 500(1 

(Manuscript accepted 6 iehn'ury J9SS) 


/\ IDt \ R, W b .inJ CiOVVI.-l- H, K. l_, 19K6. MflfllMc typc-pe-cimens 
in the Somh Australian Museum. 3. I'olyplneophoia Hec & 
)tt\t. \tti\. VW): 97 -lis, 

The South Australian Museum collection of chiton 
types is the largest ill the Southern hemisphere. It 
contains primary type material, and some secondary 
types, of 123 species, subspecies or varieties. A further 
15 species arc represented only by secondary types. The 
type status o^ at least II of the primary types is 
considered very doubtful or, in the ease of neotypes, 
invalid. I isted in an appendix are an additional 21 
species represented by types of dubious status from the 
Dupuis collection. Species are listed alphabetically 
according to the origina' name of the genus or species. 


Most o( the chiion types in the South Australian 
Museum are due to ihe early work of W. G. Torr and 
I Ashby al the turn o( t he century, particularly Ashby 
who continued his active interest in chitons well into 
the 1930$, 

Ashby donated his collection of chitons, including 
most of the types of Ion. to the South Australian 
Museum in 1932, but he continued to borrow material 
freely for his research. In 1934 Ashby's home at 
Blackwood in Ihe Adelaide Hills was destroyed by fire 
and some type material of chitons and birds was lost. 
As there are no accurate records of the material on loan 
to Ashby at the time of the fire, one must assume that 
any Ashby or Ton types, listed in the literature as being 
in the Ashby or lorr collection, and which cannot now 
be located in an Australian museum w-ere destroyed at 
that lime. 

Ashby also acquired a small collection of chitons 
from Commandant Paul Dupuis, once conehologist at 
the Museum Royal d'Histoire Naturelle de Bclgiquc, 
Brussels, including some important historical and 
probable lypc material of Blainville, D'Orbigny, Quoy 
and Gaimard, Rtn:hcbrune, Jousseaume and Dupuis. The 
Dupuis collection is not well eurated and original labels 
are often missing. Types have been indicated by a small 
label "TYPE" in red print glued to the label with the 
specimen and appears to be a later addition. As the 
type stat us of most of these specimens is very doubtful 
they have not been included in the main part of the 
following list but arc listed separately iu an appendix. 

Other chiton types in the South Australian Museum 
came from the May collection, aequired by J. C. Verco 
and presented to the museum in 1929, and as a result 
of studies by Cotton, mostly in conjunction with 
Ashby, Godfrey and Weeding. Since Cotton and 
Godfrey (1940) little of consequence has been published 
on Australian chitons and type material of only one 
species, Lucilina tiibrooki Milne, I95S (a paraiype), has 
been added to the eolleetion. 

In their monograph on South Australian chitons, 
Cotton and Godfrey (1940) and also Cotton (1964) 
selected neotypes for South Australian Bednall and 
Matthews species on ihe basis of Ashby (1918) who 
recounts the loss of Bednall and Matthews types at sea. 
However they failed to verify which types were lost as 
at least some of the original type material (syntypes) 
of Bednall is known to exist (Davis el al. 1979). 
Similarly, neotypes were also designated for other 
species where the type was presumed lost. We believe 
that in most eases these neotypes were designated in 
a somewhat arbitrary manner, the possible existence 
of type material was not sufficiently investigated and 
there were no further statements regarding the 
specimens, other than the locality and that they had 
been designated as neotypes and given museum 
registration numbers. We therefore regard all of the 
neotypes scleacd by Cotton and Godfrey (1940) and 
Cotton (1964) and listed here, with the possihle 
exception of Acanthuchiies tutci Ton & Ashby, 1898, 
as invalid. 

We I uri her believe that the South Australian 
Museum collection of chiton types is the largest in the 
southern hemisphere and one of the more significant 
collections in the world. It includes type material for 
!38 species or subspecies (excluding the Dupuis 
collection) and the type status of only II is in doubt 
or. in the ease of neotypes, considered invalid, the 
remainder have been verified by us according to 
available material and information. A further 21 
species are represented by types of dubious status in 
the Dupuis collection. 

In Ihe following list species are arranged 
alphabetically ut families under the original name at 
the time of description. Changes in familial status have 
been cross-referenced. The present status o( each 
species is, unless indicated otherwise, according to Kaas 
and van Belle (1980). The specimens are all dry except 
for sonic of the paratypes of Ischtwchiton Johnston* 

June, /V/0 


KR, S. AI.'Sl. MIS. imt &7-H5 

Jwwe. /9A0 

and /. mawsoni and are listed as "entire" when the 
articulated valves and girdle arc present or as "entire 
with animal" when the dried animal is also presenl. 
The following abbreviations arc used in the text. 
AlM = Auckland Institute and Museum, New Zealand; 
AM -The Australian Museum, Sydney; 
ANSP- Academy of Natural Sciences of Philadelphia, 
U.S.A.; BANZARR^ British, Australian and New 
Zealand Antarctic Research Expedition, 1929-31; 
MCZ = Museum of Comparative Zoology, Harvard 
University, U.S.A.; MNHN = Museum National 
D'histoire Naturelle. Paris; NMNZ = National Museum 
of New Zealand; MV = Museum of Victoria; 
N.S.W.-New South Wales; N.Z.-New Zealand; 
Qld. = Queensland; QM ^Queensland Museum; 
S.A.- South Australia; SAM = South Australian 
Museum; Tas.^Tasmania; TM-Tasmanian Museum 
and Art Gallery; W.A.= Western Australia; 
WAM= Western Australian Museum. 


Genus Acanthochites Risso, 182* 

Acanthochites cornutus Torr & Ash by, 1898 
Trans. R. Soc. S. Ausf. 22: 217, pi. 6, fig. 3a-!\ 
—Craspedochiton cornutus (Torr & Ash by, 1898). 
Holotype: D12188, 5 articulated median valves and 
associated girdle, from Marino, near Adelaide, S.A., 
at low tide, collected by E. Ashby, date of collection 
Note: Rest of type presumed lost. Type unique. 

Acanthochites crocodilus Torr & Ashby, 1898 
Tram. R. Soc, S. Aust. 22; 216, pi. 6, fig, 2a-f. 
-Notoplax crocodilus (Torr & Ashby, 1898) 
Holotype; D12137. posterior valve, one median valve 
and several fragments of valves and girdle, from 
Marino, near Adelaide, S.A., collected by W. C. Torr, 
date of collection unknown. 

Paratype: D12J95, entire specimen with animal with 
same collection data as holotype, labelled "cotype" 

Acanthochites exilis Torr & Ashby, 1898 
Trans. R. Soc. S. Aust. 22: 218, pi. 7, fig. 6a-f. 
-Craspedochiton cow urns (Torr & Ashby, 1898) 
Holotype: D12251, posterior valve and 3 disarticulated 
median valves, from Spencer Gulf or Investigator 
Strait, S.A., dredged by J. C. Verco in 10-15 fathoms, 
date of collection unknown. 
Note; Rest of type presumed lost. 

Acanthochites jucundus Rochebrune, 1882 
Bull. Soc Philom, Paris Ser, 7, 6: 194. 
=Acanthochitona jucunda (Rochebrune, 1882) 
Holotype: D12267, 1 median valve only, from New 
Holland, collected by Belligny, date of collection 

Note: According to Ashby (1926) this valve came from 
the holotype and was given to him by Dr Lamy. Recent 
enquiries indicate that the rest of the type cannot be 
located in MNHN although Ashby (1922) apparently 
saw the type and says u there are a number of specimens 
in spirit -— all much worn:' 

Acanthochites kimheri Torr, I9J2 

Trans. R. Soc. S. Aust. 36: 167, pi. 6, fig. 5a-f. 

= Acanthochitona kimheri (Torr. 1912), 

Neotypc: D13758, entire specimen with animal, from 

Aldinga, near Adelaide, S.A., collector and date of 

collection unknown. Selected by Cotton and Godfrey 

(1940: 322, fig, .515) 

Syntypes: DI2220, 3 entire specimens with animal, 

from Corny Point, Yorke Peninsula. S. A., collector and 

date of collection unknown. D12221, 2 entire specimens 

with animal and a median, posterior and anterior valve, 

from Kangaroo Island, S.A. ( collector and date of 

collection unknown. DI2227, entire specimen with 

animal, stuck cm card with other specimens from 

various localities— since removed, from Port 

Noarlunga, S.A., collector and date of collection 


Note: In the presence of syntypes a neotypc should not 
have been erected, The lypc status of 1312220 and 
particularly D12227 is very doubtful as the locality data 
do not match the original description which was 
based on 4 specimens from Aldinga and Kangaroo 
Island, However. D12220 and D1222I has the label 
"Type CO-59" which corresponds to the number of ihe 
species in the publication, indicating that these 
specimens may have been part of the original material, 
D12221 also has a label which says "cotypes type to 
be selected'*. Therefore these are almost certainly 
syntypes. The loose valves with DI222I could have 
come from the specimen illustrated by Torr and thus 
the intended type, although Torr apparently intended 
the specimen collected by Kimber from Aldinga to be 
the type. None of the above specimens are as large as 
10x4 mm which is a measurement given by Torr and 
it must be presumed that this specimen has been lost. 

Acanthochites (Lobopiax) mariae Webster, 1908 

Trans, Proc, N.Z, Inst. 40: 254, pi. 20, figs 1-11. 

^Notoplax mariae (Webster, 1908). 

Paratype: D11I08, median valve 4, girdle fragment, and 

radula, without collection data 

Note: With label "valve from Webster's paratype". 

Webster apparently did not designate paratypes 

although he mentions at least 7 specimens in addition 

to the type. Type in AIM (TM-I). 

Acanthochites matthewsi Rednali & Pilsbry, 1894 
Nautilus 1 (10): 120. 

= Notoplax (Bassethuilia) matthewsi ( Bed nail & 
Pilsbry, 1894). 



Neotype: D13741, entire specimen with animal, from 
Sultana Bay, Yorke Peninsula, S.A., collector and date 
of collection unknown. Selected by Cotton and 
Godfrey (1940: 536, fig. 534). 

Note; Iredale and Hull (1925b) selected a neotype for 
this species presuming the type to be lost (probably 
AM, C1041I). Cotton and Godfrey (1940) unaware of 
this selected yet another neolype. However, according 
to Davis el at (1979) the holotype is in ANSP (Shell 
Car, No. 64916). We therefore consider both neotypes 

Acanthochites tnaughani Torr & Ashby, I89S 
Trans. R. Soc. S. AusL 22: 218, pi. 7, fig. 5a-f. 

Acum/iachimna pilsbryi (Sykes, 1896). 
Holotype: D12264, 5 articulated median valves with 
remains of girdle and animal, plus one loose median 
valve, from Victor Harbor (Port Victor). S.A., collected 
by M M Maughan, date of collection unknown. 
Note: Rest of type presumed lost, 

Acanthochiics ruhrost rams Torr, 19|2 

Inmy R, Soc. S. Aust. 36: 169, pi. 7, fig. 7a-f. 

t Notoplax rubrostrata (Torr, 1912). 

Holotype: D13717, entire specimen with animal, from 

St Francis Island, Nuyts Archipelago, S.A., collector 

and date vf collection unknown. 

Paratype: D16020, entire specimen with animal, from 

Corny Point, Yorke Peninsula, S.A., collector and date 

ol collection unknown, 

Note: A label with the holotype says "valves figured 

pl.vji have been lost" The [ypc status of D16020 is in 

doubt as Torr (1912) only lists specimens from St 

Francis Island and Henley Beach; however a label with 

the specimen says "CO-TYPES 61" which corresponds 

to the number of the species in the publication. 

Acanthochites rttfus Torr, 1912 

Trans, /?. Soc 5. Aust. 36; 167, pi. 6, fig. 4a-f. 

= Craspedochiton variabilis (Adams & Angas, 1864). 

Holotype: DI2208, entire specimen with part of animal, 

from Kangaroo Island, S.A., collector and date of 

collection unknown. 

Note: Type unique. 

Acanthochiics subviridis Torr, 1911 

Trans. R. Soc. 8, Aust- 35: 104, pi. 25, fig. 3a-f. 

- Notoplax subviridis (Torr, 191 1 ). 

Holotype: D12872, 4 articulated median valves with 

girdle and animal, plus the 4 remaining valves 

disarticulated, from Rabbit Island, Albany, W.A., 

presumably collected by Torr, Christmas, 1910-11. 

Paratype: D14490, entire specimen with animal, with 

same collection data as holotype and labelled 

"co- type 11 . 

Acaruhovhites tatei Torr & Ashby, 1898 

Ihtns. R. Soc. 5. Aust. 22: 219, pi. 7, fig. 7a-f. 

-Acanfhochitona mmostriata (Pilsbry, 1894). 
Neotype: 1)13732, entire specimen with animal, from 
Middleton, Encounter Bay, S.A.. collector and date of 
collection unknown, Selected by Cotton and Godfrey 
(1940: 527, fig. 523). 

Note: The neotype was selected from a lot front the 
Torr collection (D12260) with the label "our type of 
Acan. tatei was very diminutive and was ruined in 
dissecting. These are from the same spot . . R 

Acanthochites tristis Rochebrunc, 1882' 

Bull. Soc. Philom. Paris Scr 7, & 194. 

— Nom. inquir. 

Holotype: D12269, I median valve only, from New 

Holland collected by Dussumier, date o\' collection 


Note: According to Ashby (1926) this valve came from 

the holotype and was given to him by Dr Lamy. Rest 

of type in MNHN 

.Acanthochites verconis Torr & Ashby, 1898 
Trans. K. Soc. S. AusL 22: 217, pi. 6, fig. 4a I. 
= Notoplax verconis (Torr & Ashby, 1898). 
Holotype: D12201, 3 articulated median valves with 
fragments of girdle and animal, plus fragments of the 
posterior valve and median valves, from Gulf Si 
Vincent, S.A., dredged by J. C. Verco, date of collection 

Note: Anterior valve presumed lost. Kaas and van Belle 
(1980) regard this species as a synonym of Notoplax 
wilsoni (Sykes, 1896) however, one of us (K.L.G.) has 
examined this species in detail and regards it a valid 

Genus Acanthochiton Gray, 1821 cm. Iredale, 1915. 

Acanthochiton berlnalli var johnsioni Ashby, 1923 
Tram, R. Soc. S. Aust, 47: 231. 
^Acanthochitona bednalfi (Pilsbry, 1894), 
Holotype: Dl 21 85, entire specimen, from Carnarvon, 
Shark Bay, W.A., collected by \V, C. Johnston, date 
of collection unknown. 

Note: Kaas and van Belle (1980) follow Cotton and 
Weeding (1939) who elevate this variety to specific rank, 
however, one of us (K.L.G.) has examined this species 
in detail and regards ii as a deep-water form of A. 

Acanthochiton brookesi Ashby, 1926 

Proc. Matac. Soc Lond, 17(1): 14, pi. I, fig. 3a-c, pi. 2, 

fig- 7 

-Acanfhochitona bmokesi Ashby, 1926. 

Holotype: DI1025, 5 disarticulated median valves and 

girdle fragments from Auckland Harbour, N.Z . 

collected by H. Suter, daic of collection unknown. 

Paratype: D1I031, entire specimen without girdle, with 

same collection data as holotype. 

Note; Remainder of holotype in AIM (TM-2). 


KK S. AIM MI'S. W t K): V7 115 

Jhw, '''• s " 

Acanthochiton gailij'fi Ashby, 1919 

Trans. R. Soc. S. Aust 43 398, pi. 42, figs 2-5, 

^Acanthochito/ui gatlifj) Ashby, 1919. 

Holotype. D12189, 2 median valves, a valve fragment, 

remains of girdle and animal, from Port Lincoln, S.A., 

collected by E. Ashby, Jan. 1917. 

Note: Rest of type presumed lost. 

Acanthochiton heterochaetus Bergenhayn, 1931 
Arkiv. Zoo/. 23A (13): 20, pi. 1, figs 38-42, pt 3, figs 

-Acanthochitona gracilis (Jeffreys, 1859). 
Syntype: DI6185, entire specimen with animal but with 
anterior valve missing, from off La Lit/., Gran Canaria, 
Canary Islands, dredged in 100 m on red algae by N, 
Odhner, 1930. 

Note: Bergcnhayn (1931) records 17 specimens but 
apparently did not dcsignaiea holotype, Our specimen 
is labelled "cotyp". 

Acanthochiton kimberi yallinuupcnsis Ashby, 1925 
Rep. Ausf. Assoc. Adv. Sci 17: 382. 
=Acanthochitona kimheri (Ton, 1912), 
Holotype: D13718, entire specimen with animal, from 
Yallingup, W.A., collected by E Ashby, 23 x.1920. 

Acanthochiton macrocyst talis Ashby, 1924 
Trans. R. Soc, S, Aust. 48: 324, pi. 31, figs 3> 3a. 
= Acanthochitona macrocystiafis Ashby, 1924. 
Holotype; D12248, 5 articulated median valves with 
remains of girdle, plus disarticulated anterior, posterior 
and median valve, from Point Puer, near Port Arthur, 
Tas., amongst holdfasts of Macrocysns pyrijera, 
collected by E. W. Mavvle, date of collection unknown. 
Paratypes: D10704, one entire specimen with same 
collection data as holotype. D12563, one entire 
specimen, one lot of 6 disarticulated median valves and 
one lot of 3 disarticulated median valves together with 
2 anterior and one posterior valves, all with the same 
collection data as holotype. 

Note; The label with D12563 indicates that the paratype 
measuring 20x9 mm was sent to Thackway. 

Acanthochiton niaxitfaris Ashby, 1919 

Trans. R. Soc. S. Aust. 43: 397, pi. 41, figs 5 & 6, pi. 

42. fig. I. 

= Acanthochitona kimheri (Torr, 1912). 

Holotype: D12253, 4 median valves and anterior valve 

artieulated with girdle, plus one loose median valve, 

Irom Marino, near Adelaide, S.A M on rocks at low tide, 

collected by E. Ashby, date of collection unknown. 

Note: Rest of type presumed lost- Type unique. 

Acanthochiton pilsbryi maughaneanus Ashby, 1919 
Trans, R. Soc. S. Aust. 43; 395, pl. 41, fig. 4. 
^Acanthochttona pilsbryi (Sykes, 1896). 
Holotype: DI5536, 3 disarticulated median \akes, 
anterior valve and valve fragments and remains of 

girdle, from Middle Harbour, Sydney, N.S.W., collected 

by I. Ashby, 6.N.1903. 

Note; Resi o\' type presumed lost. 

Acanthochiton (Notoplax) porctna Ashby, 1919 
Trans. R. Soc S. Aust. 43; 395, pl. 41. figs 7-10.. 
= Notoplax (Bassethullia) matihewsi ( ftedn all Sc 
Pilsbry, 1894). 

Holotype; D12250, specimen in two pieces with 
articulated median valves and remains of girdle and 
animal, plus disarticulated anterior, posterior and one 
median valve, from Gulf St Vincent, S.A., dredged by 
J. Ci Verco. date of collection unknown. 
Note: Type unique. 

Acanthochiton retrojectus var. pustulosis Ashby, 1922 
Trans. R, Soc. & Aust. 46; 15. 
-Acanthochitona retrojecta (Pilsbry, 1894). 
Holotype; DI2205, entire specimen with animal, from 
Quarantine Station, Sydney Harbour, N.S.W., eolleeied 
by E. Ashby, Nov. 1918. 

Acanthochiton shirlcyi Ashby, 1922 
Irons. R. Soc. S, Aust. 46; 13, pl 3, ftg, 2a-c 
= Acanthochitona shirlcyi Ashby, 1922, 

Paratype; D107I5, disarticulated valves stuck on card, 

from Northwest Reef, Capricorn Group, Qld., collector 

and date ol collection unknown. 

Note: Type in QM (MO 4043), 

Acanthochiton (Notoplax) sponxialts Ashby, 1923 
J. Roy Soc. W. Aust, 10(4); 1 3, pl. 1, fig. 1. 
- Notoplax spongialis (Ashby, 1923) 
Holotype; D13733, entire specimen from D'Emre 
casteaux Channel, southern Tas., dredged in 9-10 
fathoms by \V. L. May> date ol collection unknown. 
Note: Cottou (1964) indicated that this specimen may 
not be the holotype presumably because it does not 
match the measurements given (38 x 16 mm for D13733: 
38 X 18 mm given lor the type) and does not appear to 
match the rather poor photo of the type. However, 
SAM secrns to have the other 3 specimens mentioned 
by Ashby (DI2803 and Dl447I)and as none approach 
the dimensions of the type, and in the absence of 
evidence to the contrary, we must assume that DI3733 
is the holotype. 

Acanthochiton I hack way i Ashby, 1924 

Trans. R. Soc. S. Aust. 48. 318, pl. 31, figs I & 2. 

-Acanthochttona lhackwayi Ashby, 1924. 

Holotype: D107I6, entire specimen with animal, from 

Fly Point, Port Stephens, N.S.W., collected by E. Ashby, 

October, 1923, 

Paratype: DI6548, entire specimen with animal, from 

same locality as holotype, collected by A E, J. 

Thackway, dale of collection unknown. 

Note; Smith and Robertson (1970) list 2 paratypes in 

the MV (F16376) with the locality as Shell Harbour. 



We regard these specimens as very doubtful paratypes dredged in 60 Fathoms off Olago Heads, N.Z., by H. J. 

as Ashby only mentioned one olher specimen which Finlay, date of collection unknown. 

is the paratype in SAM. Note: Rest o\' type presumed lost. Type unique. 

Acanihoclvton turtom Ashby, 1928 

Ptvc\ Malac. Soc. land. 18(2): 79, pi. 6. figs 14, 

Acantlwchitona gamoii (Blainvillc, 1825). 
Holotype: D10985, 5 disarticulated median valves and 
girdle fragment, from Port Alfred, South Africa, 
collected by W. H, llirton, date of collection unknown. 
Paratypes: D10984, one entire specimen and two broken 
median valves, with same collection data as holotype. 
Note: The type status of some o( the valves labelled 
"holotype-' is in doubt. There are two lots. One lot of 
4 valves does not appear to belong to the type The 
valves arc from two different specimens and measure 
5.3x3.2, 5.0x3.2, 4.1x2,5 and 2.1x1.8 mm respect- 
ively. The first two valves arc too large for the type and 
the last one too small (Ashby gives a width of 4.5 mm 
for the type) and the third valve, and also the last one, 
arc green in colour and not pink as stated for the type. 
The other lot, consisting of only one valve, with the 
label "valves 6 & 7 not photo'd" would appear to 
belong to the type. 

Giles and Gosliner (1983) record another valve from 
the holotype in the South African Museum (A32649). 

Acamliochiton zealandicus doubtlcssensis Ashby, 1926 

Proc. Malac. Soc. Load 17(1): 12, pi. I, fig. 2a-c; pi. 

2, fifr 6, 

= Acanthochitona zcalandica (Quoy & Gaimard, 1835). 

Holotype: 1)11109, disarticulated specimen, from 

Doubtless Bay, N.X., collected by A, L\ Brookes, date 

of collection unknown. 

Paratypes: Dill 18, 3 entire specimens, with same 

collection data as holotype. 

Note: Paratypes correspond to Ashby's paratypes 

No. 1, 3 and 4. 

Genus Cruspcdodiifnn Shutticworth, 1853 

Craspcdochiton jaubertensis Ashby, 1924 
Trans. R. Soc. S. Aust. 48: 326, pf. 31, tig. 5a-c. 
Holotype: D1I234, 2 articulated median valves with 
fragments o\^ one valve and remains of girdle and 
animal, plus disarticulated anterior, posterior and 
remaining 3 median valves, dredged in 70 ft, 42 miles 
W.S.W. of Cape Jaubert, northern W.A., by Dt h. 
Mjoberg, 26.V.I91I. (Swedish Scientific Expeditions 

Genus Ijtphoplax Ashby, 1926 

Lophop/ax jinlayi Ashby, 1926 

Proc. Malac Soc. Lond, 17(1): 30, pl. 3, fig. 4, pi. 4, 

fi&l 1-4. 

-Craspedochnon rubi^inosus (Huttoit, 1872). 

Holotype: D143I8, 5 disarticulated median valves. 

Genus Notoplax Adams, 1861 

Notoplax (Amblyplax) brookesi Ashby, 1929 

Trans. Proc. V.Z. Inst. 60: 370, pi 32, litis 1-4. 

= Notoplax brookesi Ashby, 1929. 

Holotype: D1U)30, 2 disarticulated median valves, fiom 

fauranga Harbour, N.Z., dredged in 3 fathoms, 

collector and date of collection unknown. 

Paratype: DIIJB, entire specimen with same collection 

data as holotype. 

Note: Rest of type presumed lost. 

Notoplax (Amblyplax) foveauxensis Ashby, 1926 
Proc, Malac. Soc. I ond. 17(1): 20 pl. I, fig. 5a-c 

Craspedochtton rubtgtnosus (Hnlton, 1872). 
Holotype: D16021, I median valve, from FtoV&aW 
Strait, N.Z., dredged in 15 fathoms on oyster shell, 
collected by W. R. B. Oliver, date of collection 

Paratype: DI6022, 4 articulated median valves witb 
remains a( girdle and animal, plus disarticulated 
anterior, posterior and 2 median valves, with same 
collection data as holotype. 
Note: Rest of type in NMNZ (Ml 584). 

Notoplax (Amblyplax) mariae haurakiensis Ashby, 


Proc. Mulac. Soc. fond. 17(1): 26, pl 2, fig. 3a-c. 

Notoplax nutria? (Webster, 1908). 
Holotype: Dl 1043, 4 articulated median valves and 
remains of girdle, plus one disarticulated median valve, 
from Hauraki Gulf, N.Z., dredged in 20 fathoms, off 
Atrtna shell, collected by A. E. Brookes, dale of 
collection unknown. 

Paratype: D16550, remains of girdle, stuck on card, 
wiLh same collection data as holotype. 
Note: Rest of type in AIM (fM 535). 

Notoplax (Amblyplax) oliveri Ashby, 1926 

Proc. Malac. Soc. Lond. 17(1): 18, pl. 1, fig. 4a-c. 

-Craspedochtton rubtginosus (Hutton, 1S72). 

Holotype: D11044, 1 median valve, from between 

Kawau and Tiritin islands, Hauraki Gulf, near 

Auckland N.Z., dredged in 20 fathoms in dead Atrina 

shell, collected by A. li. Brookes, date of collection 


Paratype: D16551, 5 articulated median valves with 

remains of girdle, plus disarticulated anterior, posterior 

and median valve, with same collection data as 


Note: Rest of type in NMNZ (M1585). Paraiype has 

label "Ashby's No. 2" 

Notoplax rottnestensis Ashby, 1929 

./. Roy. Soc. IT Aust. 15: 47, figs 10-13- 


Kl-( S AUST NIL'S 19(8): 97-115 

.ftitw, 19X6 

Holotype: DI2565, 3 disarticulated median valves (2, 
3, & 7), fragments of girdle and raduta mounted on 
slide, From Bathurst Point, Rottnest island, W.A., 
collected by L. Glauert, date of collection unknown. 
Note: Rest of type in WAM (12885). 


Genus Catlistochiton Dall, 1882 

Callistochiton antiquus mayi Ashby, 1919 
Trans* *t Soc. S. Aust, 43: 401, pi. 42, figs 8 & 9. 
-Cullistochifon antiquus mendionalis Ashby, 1919. 
Holotype: D12550, 1 median valve from Penguin rocks, 
north-western Tas., collected by E, Ashby. M.x.1916. 
Note: Rest of type presumed lost. 

Callistochiton antiquus meridionalis Ashby, 1919 
Trans. R. Sot. S. Aust 43; 400, pi. 42. fig. 7. 
Holotype: D13716, disarticulated valves and piece of 
girdle and radula, from Marino, near Adelaide, S.A., 
collected by E. Ashby, date of collection unknown. 

Ischnochtton (Lepidozona) asfhenes Berry, 1919 
^Callistochiton asthenes (Berry, 1919). 

Callistochiion augustensis Ashby # Cotton, 1937 
Trans. R. Soc. S. Aust. 61: 145, pi. 8, figs 2-4, 
Holotype: D12952. anterior valve, 6 disarticulated 
median valves with one broken and remains of animal, 
from Fori George IV, Augustus Island, northern W.A., 
collected by B. Bardwcll, October, 1933. 
Note; According to Ashby and Cottoa(1937) there were 
only six detached valves so presumably the posterior 
valve was lost before the description of the species. Type 

Callistochiion broomensis Ashby & Cotton. 1934 
./. Roy Soc U[ Aust. 20: 213, pi. 13, fig. 3. 
Holotype: DI0723, entire specimen with animal, from 
Ganihcaume Point, Broome, W.A., collected by MCZ 
expedition to Australia, September, 1929. 
Note: Type unique. 

Callistochiion clenchi Ashby & Cotton, 1934 
J. Roy. Soc. W. Aust. 20: 214, pi, 13, fig. I. 
Holotype: D10724, entire specimen with animal, from 
Gamhcaume Point, Broome, W.A., collected by MCZ 
expedition to Australia, September, 1929. 
Note: Type unique. 

Callistochiion ntawlei Iredale &. May, 1916 
Proc. Malac. Soc. Lond. 12(2/3): 113, pi. 4, Fig. 5. 
Paratypes: D120I9, one disarticulated specimen with 
dried radula, locality not indicated bui probably Tas,, 
collector and date of collection unknown; D12026. 
disarticulated valves of one specimen with same 
collection data as Dl 201*9. 

Note: An old label with the specimens says that they 
were disarticulated by Iredale and used in the original 
type description but there is no indication that they 
were designated paratypes. Types id TM (E 201/7542). 

Callistochiion occidus Ashby &. Cot l on, 1934 
J. Roy. Soc, U\ Aust. 20: 215, pi. L3, fig. 4. 
Holotype: D10725, entire specimen with animal, from 
Gantheaume Point, Broome, W.A., collected by MCZ 
expedition to Australia, September, 1929. 
Note: Type unique. 

Genus Lophochiton Ashby, 1923 

Lophochiton johnstoni Ashby, 1923 

Tmns. R. Soc, 5. Aust. 47: 234, pi. 16. fig 7a-c; pi. 17. 

fig. la-d. 

= Callistochiion coccus (Menke, 1844). 

Holotype: D12I52, disarticulated specimen with one 

median valve fragmented, dried radula and piece of 

girdle, from Carnarvon, in extreme north of Shark Bay, 

W.A., collected by W. C, Johnston, date of collection 


Note: Type unique. 


Genus Acutoplax Cotton & Weeding, 1939 

Acutoplax cottom Weeding, 1940 
Trans. R. Soc. S. Aust. 64(1): 48, pi. 4, fig. I, la. 
= Callochiton cottoni (Weeding. 1940). 
Holotype: D13766, entire specimen with animal, 
dredged in shallow water, Spencer Gulf, S.A., by 
Fisheries launch "WhyaJla", K, Sheard. March, 1938. 
Note: P, Kaas, Rijksmuseum, Leiden, has examined the 
type and considers it a synonym of Callochiton mayi 
Torr, 1912. 

Genus Callochiton Gray, 1847 

Callochiton elongatus May, 1919 
Pap. Proc. R. Soc. Tas. 1919: 55, pi. 14, fig. la-b. 
Paratypes: D12696, 7 entire specimens (3 with animal), 
from Norfolk Bay, Tas., collected by E. Mawle, 1915. 
DI0681, 6 entire specimens stuck on card, 2 from Port 
Arthur, Tas., collector and date of collection unknown, 
and 4 from Woodbridgc, las., collected by E. Ashby, 

Note: D12696 are from the May collection (No. 230) 
and are labelled "cotypes". DI0681 arc from the Ashby 
collection and are also labelled "cotype" probably 
referring to the two specimens from Pori Arthur, 
however May notes only 7-8 specimens so the type 
status of these specimens is in doubt. The 4 specimens 
from Woodbridge cannot be types- Type in TM (El 77/ 



Calloclvton klemi Ashby, 1926 yncl 4 median valves, from Maud's Landing, north of 

Trans, R Soc S. Aitsr. 50: 243, fig. 4. Shark Bay, northern W. A., collected by T. Cutton, date 

Holotype: Dl 1703, 1 median valve, from amongst shell of colleciion unknown. 

grit, Daly Head, Yorke Peninsula, S.A., collected by Note: Median valves 2 and 6 presumed lost. 

W. Klem, dale of collection unknown, 

Nuic; lype description was based on this single valve. Genus Chiton Linnaeus, 1758 

Callochiton platessa van fossa Ashby, 1922 
Trans. R. Soc. S. Aim, 46: 19, pi. 3, fig. 4. 
= Callochiton crocinus (fiCCVfc, 1847) 
Syntypes: 011699, 3 entire specimens (2 with animal 
remains), from WatsoiVs Bay, Port Jackson, N.S.W., 
collector and date of colleciion unknown. 
Note- The above three specimens were sent to Ashby 
by May and Ashby refers to them in his description 
and illustrated the larger specimen. The other 
specimens mentioned by Ashby do not appear to be 
in SAM although one specimen, D11694, with the label 
"variety with pits on valve T could be the SAM 
specimen mentioned by Ashby. Another specimen in 
SAM, U10071, is labelled "Holotype" but it does not 
match Ashby's description as the pits on valve 7 are 
poorly defined and only 6-7 in number. It is stuck on 
a card with 3 other specimens and is the only one 
labelled "var fossa" and we suspect that this 
information was added after the publication of the 
original description. 

Callochiton rufus Ashby. 1900 
trans. R. Soc, S. Aust. 24: 87, pi. 

L I'g 



Holotype: DII700, specimen with first median valve 
removed and anterior valve missing, presumed lost, 
with animal, from Gull St Vincent, S.A., dredged by 
.1. C. Verco, date of collection unknown. 
Note: Type unique. 

family CH1TOMOAE 

C.enus Acanthopleura Guilding, 1829 

Acanthop/eura gemmata var. queenslandicu Ashby, 


J. Roy. Soc. It Aust. 8: 30. 

\<vnthopleura gemmafa (Blainville, 1825). 
Holotype; 012459, disarticulated specimen with girdle 
fragments, from Dunk Island, Qld., collected by J, 
Shirley, date of collection unknown. 
Note: There is also a radula, mounted on a slide, 
labelled "Acanthopleura gemnuua Bl. var 1, Dunk Id. 
Q" which could have come from the above specimen. 
This same specimen W'as selected by Ashby (1928) as 
the neotypc of Chiton gemmatus Blainville, 1825 (also 
listed here). 

Acanthopleura gemmatus maudensis Ashby, 1928 
Trims. R. Soc, S, Aust, 52; 172, pi. 12, figs 8 & 9. 
^Acanthopleura gemnwta (Blainville, 1825). 
Holotype: D10775, disarticulated anterior, posterior 

Chiton aureomaculata Bednall & Matthews, 1906 
Proc. Matac. Soc. Lotl 7(2): 91, pi. 9, figs 3 & 3a- 1, 
^Chiton (Rhyssoplax) tricostalis Pilsbry, 1894. 
Neotypc: D14135, entire specimen with animal, from 
Cape Banks, S.A., collector and date oi collection 
unknown. Selected by Cotton (1964: 94). 
Note: From Torr collection. Listed by Cotton (1964) 
as neotypc without discussion and considered, by us, 
as invalid. 

Chiton hednalli Pilsbry, 1395 

Nautilus 9: 90. 

^Chiton (Rhyssoplax) hednalli Pilsbry, 1895. 

Holotype: 013742, disarticulated specimen with 

remains of animal, from Sultana Bay, Yorke Peninsula, 

S.A., collected by W. T. Bednall, dale of collection 


Note: From Bednall collection. 

Chiton exoptandm > Bednall, 1897 
Proc. Malac. Soc. Lo/uL 2(4): 152. 
= Chiton (Rhyssoplax} exoptandus Bednall, 1897. 
Neotypc: D13744, entire specimen, from Sultana Bay, 
Yorke Peninsula, S.A., collector and date o\ collection 
unknown. Selected by Cotton and Godfrey (1940: 559). 
Note: Listed by Cotton and Godfrey (1940) as neotypc 
without discussion and considered, by us, as invalid. 

Chiton gemmatus Blainville, 1825 
Diet. Sci. Nat. 36: 544. 

= Acanthopteura gemmata (Blainville, 1825).. 
Ncotype: D12459, the same specimen as the holotype 
of Acanthopteura gemruafa var queenslandicu Ashby, 
1921, also listed here, Selected and figured by Ashby 
(1928; 172, fig. 6 & 7). 

Chiton (Rhyssoplax) kimberi Ashby, 1928 
Trans. R. Soc. S. Aust. 52; 170, pi. 12, figs 10-12, 
Holotype: 012393, disarticulated anterior, posterior 
and 3 median valves plus fragments of girdle and radula 
mounted on slide, from Capricorn Group, Qld., 
collected by W. J. Kimber, date of collection unknown- 
Note: Rest of type presumed lost. Type unique. 

Chiton marmoreus var. coeruleus AVinkley, 1894 

Nautilus 8(4): 78. 

^Tonicella marmorea (Facbicus, 1780) (1SCHNO- 


Paratypc: D10467, entire specimen, from Eastport, 

Maine, U.S.A. , collected by H. W, Winkley, date of 

collection unknown. 


KLC. S. AUSI. MUS. \9{H): 97-115 

JtWi\ /VAV> 

Note: Specimen with original MCZ label and with the 
number "1707" written on the shell. Horn Ashby 
collection. Type in MCZ (MCZ 32856). 

Chiton oruktus Maughan, 1900 

Trans. R. Soc 5. AMI* 24: 89, pi. 1, fig. 3a-g. 

^Chiton (Rhyssoplax) oruktus Maughan, 1900. 

Holotype: 1)1489, disarticulated specimen, plus girdle 

fragment, from MacDonnell Bay, S.A., collected by 

W. G. Torr, date of collection unknown. 

Paratype: D12383, entire specimen, from Robe, S.A., 

collector and date of collection unknown. 

Note: The type status of D12383 is doubtful as the only 

locality listed by Maughan (1900) was MacDonnell Bay. 

Chiton scaber Blainvjlle, 1825 
Did. Sci. Nat. 36: 553. 
= Norn, inquir. 

Holotype: D12271, I median valve only, from New 
Holland, collector and date of collection not indicated. 
Note: From Ashby collection with label "valve of type" 
presumably obtained by Ashby during his visit to 
Europe in 1922, However, the species is not mentioned 
by Ashby (1922). 

Chiron tulipa alfredensis Ashby, 1928 

Proc Maiac. Soc. 18(2)1 87, pi. 8, figs 19-21. 

- Chiton (Rhyssoplux) tulipa Quoy & Gaimard, 1835. 

Holotype: D10983, 3 disarticulated median valves, 

girdle fragments and loose girdle scales, from Port 

Alfred, South Africa, collected by W. H. Turton, date 

of collection unknown. 

Paratypcs: 011157, 3 entire specimens with same 

collection data as holotype. 

Note: Holotype with label "these valves not photo'd" 

Rest of type presumed lost. 

Chiton verconis Torr & Ashby, 1898 
Trans. R. Soc & Aust. 22: 215, pi. 6, fig. la-f, 
- Chiton (Mucrosc/uama) verconis Torr & Ashby, 1898. 
Lectotypc: D16546, entire specimen with animal, from 
Gulf St Vincent, S.A., dredged by W. J. Kimber, date 
of collection unknown. (Leclotype chosen here). 
Paralectotype: DI6547, entire specimen with animal, 
with same collection data as leclotype. 
Note: The SAM register and Cotton (1964) list D12380 
as the holotype but this lot consists of one complete 
specimen, one median valve and the dried remains of 
a specimen from which the valves have been removed, 
none of which corresponds to the original description 
and is not even C. verconis but Callochiton cottoni 
(Weeding, 1940). In view of this confusion we have 
selected the above lectotypc and paralectotype from a 
lot which we believe constituted syntype material. The 
lectotypc (Fig. 1A) matches the original description and 
measures 17x11 mm (curled) and 31 mm along the 
curled dorsal surface, which approximates to the 
measurement of 24x 12 mm given by Torr and Ashby, 
and may have been the specimen for which this 
measurement was given. The paralectotype (Fig. IB) 

MG, I. Chiton verconis lorr <V A->hb\. IS9S. A. Leaotype, 17> tl mm (curled), \\. Parcik\Moi\pc, 11.5*6.0 mm. 



\\ hich was collected wiih the leclotype is less curled and 

measures 11.5x6.6 mm. 

Another 3 specimens (DI0739, DI2379 and D12674) all 

dredged in Gulf St Vincent by Verco may also have 

belonged to the original type series bul they are not 

given type status by us. 

The valves figured by Torr and Ashby are presumed 


Genus liolophura Pilsbry, 1893 

Plaxiphora pustulosa Torr, 1911 

- I mfophura (Cluvarizonu) hirtosa (Blainville, 1825). 

Genus Lucilina Dall, 1882 

Lucdtnu tainfordiana Hull, 1924 

Proc. R. Soc. Old. 36: 115, pi. 21, fig. 3. 

-ionicia (Lucilina) fortdirata (Reeve, 1847). 

Paratype: 1)14496, entire specimen with animal, from 

North Head, Port Denison, Qld, collected by E. H. 

Rainford and A. F. B. Hull, 18.ix.l923. 

Note: From Hull collection. Type in QM (MO 1139). 

LucUina idhrooki Milnc f 1958 

Pmc, R. Zool. Soc. N.S.W. 1956-57: 152, fig. 1-4. 

= Tonicia (Lucilina) tilbrooki (Milne, 1958). 

Paratype: D14591, entire specimen with animal remains, 

from Heron Island, Qld., collected by K. L. Milne, 


Note; Type in MV (FI847I). 

Genus Muerosquama Iredale & Hull, 1926 

\lui-m\qutimu nieiseni Cotton & Weeding, 1939 
Trans. R. Sot. S. Aust. 63(2): 190, pt. 7, fig. 8. 
= Chiton (Muerosquama) curnosus Angas, 1867. 
Holotype: D13720, entire specimen, from Hardwicke 
Bay. Yorke Peninsula, S.A., dredged by J. C. Verco, date 
of collection unknown. 
Note: Type unique. 

Muerosquama sheardi Cotton & Weeding, 1939 
Trans. R. Sat; & Aust. 63(2): 190, pi. 7, fig. 3. 

— Chiton (Muerosquama) verconis Torr & Ashby, 1898. 
Holotype: 1)13721, entire specimen with animal, from 
Spencer Gulf, S.A., dredged by Fisheries Launch 
"Whyalla", K. Sheard, March, 1938. 

Note: From Ashby collection. Listed by Cotton and 
Godfrey (1940) as neotypc without discussion and 
considered, by us, as invalid. 

Onithochiton quercinus occiden talis Ashby, 1929 

Trans, R. Sac S, Aust, 53; 65, 

Holotype: D12527, entire specimen, from Dongarra, 

W.A., from exposed outer reef, collected by E. Ashby, 


Genus Rhyssoplax I hiele, 1893 

Rhyssoplax jacksonensis Ashby, 1921 

Proc. R. Soc. Vic. (NS) 33: 153, pi. 8, fig. la~b. 

= Chiton (Muerosquama) carnosus Angas, 1867, 

Holotype: D10720, entire specimen, from Quarantine 

Station, Port Jackson, N.S.W., collected by E. Ashby, 


Rhyssoplax surrecta Hull, 1922 
Aust. Zooi 2(3): 84, pi, 248. figs 1-4. 
-Chiton (Rhyssoplax) bednalii Pilsbry, 1895. 
Holotype: D1267I, entire .specimen, from Port 
Willunga, near Adelaide, S.A., collected by W. I. 
Kimbcr, date of collection unknown. 
Nole; Type unique. 

Genus Sypharochiton Thicle, 1893 

Sypharochiton /tellisserpenfis scptentriones Ashby, 


Trans. R. Soc, S. Aust. 48: 321. 

= Chiton pellisserpentis Quoy & Gaimard, 1835. 

Holotype; DI7238, entire specimen with animal, from 

Nelson Bay, Port Stephens, N.S.W., collected by H. W. 

Thackway, October, 1923. 

Genus Tonicia Gray, 1847 

Tonicia htdlianus Torr, 1911 

Trans. R. Soc. S. Aust. 35: 104, pi. 25, figs 4a-f. 

=Tom'cia (Lucilina) huliiana Torr, 1911. 

Holotype: DI2873, entire specimen from Ellenbrook, 

south of Cape Naturaliste, W.A., collected by W. G, 

Torr, Christmas, 1910-11. 

Note: Type unique. 

Genus Choriplax Pilsbry, 1894 

Genus Onithochiton Gray, 1847 Choriplax grayi pattisoni Ashby, 1921 

Trans. R. Soc. S. Aust. 45: 137, pi. 9, fig. la-c. 

Ondhochiton ushbyi Hcdnall & Matthews, 1906 Holotype: D15019, entire specimen, found with giant 

Proc. Maiac. Soc. Lon. 7(2): 92, pi. 9, figs 2; 2a-e. kelp {Laminaria) washed ashore near Cape Banks 

Neotype: D13745, entire specimen with animal, from Lighthouse, S.A., collected by G. Pattison, 1921. 

south of Port Willunga jetty, near Adelaide, S.A., Note: One ol us (K.L.G.) has examined this species in 

collected by E. Ashby, 2.iv t 1926. Selected by Cotton and detail and regards it a synonym of Choriplax gmvi 

Godfrey (1940: 566). (Adams & Angas, 1864). Type unique. 


KLC. S. AUST. MUS. 19(8): 97-115 

June. fPfr 


Genus Cryptoplax Blainville, 1825 

Cryptoplax iredalei Ashby, 1923 
Trans, R, Soc S, Aust, 47; 238, pi. 19, fig. 4, 
Holotype: D12306, entire specimen, from Port Lincoln, 
collected by E. Ashby, January, 1917, 

Cryptoplax si rial us Van wesfenwnsis \shby\ 1923 
Trans. R. Soc, S, Aust. 47: 238. 
- Cryptoplax striata westernensis Ashby, 1923. 
Holotype; D10717, entire specimen with animal 
remains, from Rottncst Island, W.A., collected by E. 
Ashby, 2.x, 1920. 
Note: Type unique. 


Genus Anisoradsia Iredale & May, 1916 

Anisoradsia mawlei saundersi Ashby, 1918 
Trans. R. Soc. & Aust. 42.* 82. 
—Ischnochiton (Hererozona) cariosus Pilsbry, 1892, 
Holotype: D11961, specimen with anterior, posterior 
and 1 median valve disarticulated, from Port Lincoln, 
S.A., collected by E. Ashby, January, 1917. 

Genus Dinoplax Dal I, 1882 

Dinoplax gigas var. vatidjossus Ashby, 1934 

Ann, Durban Mus. 3(4): 79, pi. 9, fig. 3. 

= Dinoplax validfossus Ashby, 1934. 

Holotype: D10971, entire specimen, from Durban, 

Natal, South Africa, collector and dale of collection 


Note: From J. D. Casey collection. 

Genus Ischnochiton Gray, 1847 

Ischnochiton ft.epidozona) asthenes Berry, 1919 

Lorquitua 2(6): 47, 

^Cailistochiton asthencs (Berry, 1919) (CALLISTO- 


Paratype; D10404. entire specimen, from White's Point, 

Los Angeles County, California, U.S.A., collected by 

A. G. Smith, 14-I8.vii.1916. 

Note: Label with specimen has the number ,( SSB 1196". 

Type in Berry Collection. 

Ischnochiton atkinsoni Jrcdalc & May, 1916 

Proc. Ma/ac Soc. ton, 12(2/3): 110, pi. 4, fig. 3. 

Paratypes; D15678, 8 entire specimens with animals, 

from Stanley to Devonport, Tas., collected by W. G. 

Torr, date of collection unknown. 

Note; From May collection (No. 233). Type in TM 


Ischnochiton atkinsoni hruniensis Ashby, 1927 
Pap. Proc. R. Soc. Tas. 1926: 111. 

Holotype; D1I966, entire specimen, from Limawanna, 
South Bruny Island, D'Enlrecasteaux Channel, Tas., 

collector and date of collection unknown. 

Ischnochiton atkinsoni lincolnensis Ashby, 1920 
Trans. R, Soc. S. Aust. 4: 275, pi. 12, fig. 5a-b. 
^Ischnochiton variegatus (Adams & Angus, 1864). 
Holotype: Dl 1763, entire specimen, from Port Lincoln, 
S.A., collected by E. Ashby, January, 1917. 

Ischnochiton aural us Ashby, 1920 

Trans, R, Soc. S. Aust, 44: 277, pi. 12. fig. 6a-b. 

—Ischnochiton variegatus (Adams & Angas, 1864). 

Holotype: D11850, entire specimen with animal, from 

Marino, near Adelaide, S.A., collected by E. Ashby, 

date of collection unknown. 

Paratype: D10738, entire specimen with animal, with 

same collection data as holoiype. 

Ischnochiton hakeri Torr, 1912 
Trans. R. Soc. S. Aust. 36: 169, pi. 7, fig. 8a-c, f. 
= Notti, inquir. 

Holotype: D12145, entire specimen with animal but 
posterior valve missing, from Henley Beach, Adelaide, 
S.A., collected by W, H. Baker, date o^ collection 

Note; The specimen is very eroded and the posterior 
valve was missing at the time of collection. Type unique. 

Ischnochiton (Isochiton) bardwel/i Ashby & Cotton, 


./ R. Soc. IV. Am, 20: 217, pi. 13, fig. 5. 

^Ischnochiton bardwelli Ashby & Cotton, 1934. 

Holotype: 1)11978, disarlieulaicd specimen and girdle 

scales mounted on slide, from off Broome, VV.A,, 

dredged in 7 fathoms by B. Barclwcll, date of collection 


Paratypes; DI1979, 2 entire specimens wifh same 

collection data as holotvpe, 

Note: Smith and Robertson (1970) list the holotype as 

being in the MV (F17977). However, our specimens 

match ihe original description and are clearly marked 

"holotype" and "paratypes". Also we have the original 

label u Solvaga recens Th\e\e" as mentioned by Ashby 

and Cotton, so that the type status of the MV specimen 

must be very doubtful. 

Ischnochiton bednalli Torr, 1912 

Trans. R. S. Aust. 36: 166, pi. 5, fig. 3a-f. 

-Subterenochiton bednalli (Torr, 1912) 


Holotype: D11792, entire specimen and radula mounted 

on slide, from St Francis Island, Nuyts Archipelago, 

S.A., collector and date of collection unknown. 

fscimochiion (Hupioplax) broonwnsis Ashby & 

Cotton, 1934 

J, R, Soc. U: Aust, 20: 216, pi. 13, fig. 2. 



Holotypc; D10729, disarticulated specimen with radula 
and girdle remains mounted on slides, from 
Gantheaume Point, Broome, W,A„ collected by MCZ 
expedition to Australia, September, 1929. 
Paratype: D15537, entire specimen with animal, with 
same collection data as holotype. 

Ischnochiton (Heterozonaj curiasvs var. occidentalis 

Ash by, 1921 

Trans. R. Soc. S. Aust. 45; 42. 

- Ischnochiton ( Heterozonaj cariosus Pilsbry, 1892. 

Holotype: DU884, entire specimen with animal, from 

Point Peron, near Rockingham. W.A., collected by E. 

Ashby, 1920. 

ischnochiton (Radstella) delugouensis Ashby, 1931 
,4/;/? S. Afr, Mus. 30(1)! 40, pi. 6, figs 63-66. 
^Ischnochiton defagoaensib Ashby, I93L 
Holotype; D16019, girdle fragments mounted on slide 
only, from Delagoa Bay, South Africa, collected by 
K. H. Barnard, date of collection unknown. 
Note; Rest of type in South African Museum (A 6589), 
Type unique. 

Ischnochiton hewitti Ashby, 1931 

Ann, & Aft Mus. 30(1)- 33, pi. $[ figs 50-53. 

=-Ischnochiton bergoti (Velain, 1877). 

Holotype: D 1 0997, entire specimen and radula 

mounted on slide from Table Bay, South Africa, 

collector and date of collection unknown. 

Note: The above specimen is labelled "type" and with 

the South African Museum No, 4 '6757". However, 

Ashby (1931) figured the specimen from the Albany 

Museum (No. 8085) as the type. The above specimen 

therefore cannot be the type but is probably a paratype. 

Giles and Gosliner (1983) are similarly in error as they 

list a valve from the type, also numbered A 6757. 

Ischnochiton ireclafei Dupuis, 1918 
Bull, Mus. Nat Hist. Nat. 24(7): 526. 
^Ischnochiton lineolatus (Blainvillc, 1825). 
Syntypes: D12718, 7 entire specimens, 6 from Penguin 
and one from Stanley, north -western Tas., collector and 
date of collection unknown. 

Note: From May collection (No, 236) labelled "para- 
types", [he type status of these specimens is very 
doubtful as Dupuis (1918) did not give specific details 
of specimens and says that he did not sec any local 

Ischnochiton iredalei kingensis Ashby &, Hull, 1923 
Aust, ZooL 3(2): 81, pK 8, figs 1-4. 
-Ischnochiton hneolarus (Blainville, 1825). 
Holotype: D13731, median valves 3-5 and parts of 
girdle stuck on card, remaining valves disarticulated, 
from Fraser Bay, King Island, Bass Strait, collected by 
A. F. B. Hull, December, 1922. 

Ishnochiton (sic.) jervisensis Ashby & Cotton, 1937 
Trans. /?, Sac. S, Ausf. 61(1): 147, pi. 8, fig, 1. 
= Ischnochiton pilsbryi Bednail, 1897. 
Holotype: D13282, entire specimen, from Cape Jcrvis, 
S.A., in a sheltered pool at low tide, collected by F, 
Ashby, 28. i. 1937. 
Note: Type unique? 

Ischnochiton johnstoni Cotton 1937 
BANZARE Rep Set; B. 4(1): II, figs 10-18. 
Holotype; DI4456, disarticulated valves stuck on glass 
and radula mounted on slide, trom BANZARE Stn. 
47 (49-50 S, 63-33 'H), off Kcrguelcn Island, depth 
150 rn, collected by BANZARE, 7.ii.l930. 
Paratypes: DJ4456, disarticulated valves of one 
specimen stuck on glass with holotype and radula 
mounted on slide, with same collection data as 
holotypc. D15162, 2 entire specimens with animal, with 
same collection data as holotype, DI60I5, 3 entire 
specimens with animals, in spirit, with same collection 
data as holotype. 

Ischnochiton Levis Torr, 19t2 
Trans. R. Soc. & Aust 36- 168, pi. 6, fig. 6a-f. 
^Ischnochiton (Autochiton) levis Torr, 1912. 
Holotype: D11976, 5 articulated median valves with 
girdle and animal and disarticulated posterior valve, 
from Edithburgh, Vorke Peninsula, S.A., collected by 
E. H. Matthews, date of collection unknown. 
Note: Rest of type presumed lost. Type unique. 

Ischnochiton (AnisoradsiaJ mawlei Iredaje & May, 1916 
Proc. Malac. Soc. Lond. 12(2/3): 108, pi. 4, fig. 4. 
-Ischnochiton mawlei lredale & May, 19J6. 
Paratype: DI2546, disarticulated specimen and remains 
o\ girdle, from South Tas., collector and date of 
collection unknown. 

Note: A label with the specimen says (i dissected by 
Irc'dale", a later SAM label says "Holotype (?)". The 
type does not appear to have been disarticulated, 
according to the original description, and is located in 
TM (E 196/7537) thus the SAM specimen is most likely 
a paratype, 

Ischnochiton mawsoni Cotton, 1937 
BANZARE Rep. Set: B, 4(1): 9, figs 1-9. 
Holotype: D14457, disarticulated valves stuck on glass 
and radula and girdle mounted on slide, from 
BANZARE Stn. 83 (54°42'30"S, 158°54'30"E), off 
Lusitania Bay, Macquarie Island, depth 69 m, collected 
by BANZARE, 5.xiiJ930. 

Paratypes: D15160, about 300 specimens with animals, 
with same collection data as holotype. DI5J6I 
disarticulated valves of one specimen stuck on glass 
with holotype and radula and girdle mounted on slide, 
with same collection data as holoupe. D16014, 55 entire- 
specimens with animals, in spirit, with same collection 
data as holotype. 


RD S \LM. Ml S l'^S): M7-II5 

./•Wffi /1AV6 

Ischnochiton (Haploplax) mayi van v/V/r/As Ashby. 1920 
Trans. R. Soc. S. Aust. 44: 264. 
= Ischnochiton (Haploplax) muyi Pilsbry, 1895. 
Holotype: D11972. entire specimen with animal, from 
Lunawanna, Soulh Bruny Island, D'LntTecasteaux 
Channel, Tas., collected by fc. Ashby, 22.iii.1920. 

Ischnochiton (Haploplax) tnisintaensis Ashby, 1 923 
Ttvns. R. Soc. S. Aust. 47: 228, pi. 16, figs 6, 6a-c. 
^Ischnochiton (Haplopiax) adelaidensis (Reeve, 1847). 
Holotype: D12498, disarticulated specimen and 
fragments of girdle, from Misima, Papua New Guinea, 
collected by R. Andrew, date of collection unknown, 
Paratype; D16552, entire specimen with animal, with 
same collection data as holotype. 

Ischnochiton (Stenochiion) pa/lens Ashby. 1900 
Trans. R. Soc. S. AttsL 24: 86, pi. 1, fig, la ft $ 
^Stcnochiton pollens (Ashby, 1900). 
Holotype: D978, 5 disarticulated median valves and the 
anterior valve, from Gulf St Vincent, S.A., dredged by 
J. C. Verco, dale of collection unknown. 
Paratype: D11728, entire specimen with animal, with 
same collection data as holotype (labelled 4, cotype"). 
Note: Rest of type presumed lost. 

Ischnochiton (Stenochiion) pilshryunus Bednall, 1897 
Prat. Malac. Soc. Land, 2(4): 142. 
^ Stenochiion pilsbryanus (Bednall, 1897). 
Neotype: D11729, posterior and 3 median vylvt _s 
articulated with remains of girdle and disarticulated 
anterior and median valve, from Tapley Shoal, Gulf Si 
Vincent, S.A., on Zosterahcd, collected by R. Tate, date 
of collection unknown. Selected by Ashby (1919: 67 ? 
pi. 11, figs 2, 2a-e). 

Paraneotype: Dl 1727, entire specimen from Marino, 
near Adelaide, S.A., collected by E. Ashby, I9.ii.19l0. 
Note: The above specimens are the "holotype" and 
"paraiype" respectively of Ashby *.s (1919) description 
of Stenochiion (Zosiericola) pilsbryanus (Bednall, 
1897) According to Davis el ai (1979) there arc at least 
3 synlypes of this species in ANSP (Shell Cat. No. 
69142) however, Ashby (1919) refers to this material and 
says "I easily identified in the material shown to me 
the three species S jutoides, Ad. and Ang.; S. 
cymodoceulis, A.shby; and S. posidoniafis, Ashby, all 
very small and juvenile". Later, Ashby (1927) doubted 
I he validity of his identification o\ the neotype and 
renamed it as a new species, Stcnochiton tatei, also 
listed here. 
Resl of neotype presumed lost. 

Ischnochiton pilshryt Bednall, 1897 
Proc. Malac. Soc. Lond. 2(4): 143, pi. 12, figs 2, 2a-e. 
Neotype: DI1766, entire specimen, from Sullana Bay, 
Yorkc Peninsula, S.A M collector and dale of collection 
unknown. Selected by Cotton and Godfrey U940: 491, 
fig 477), 

Note: From Matthews and Bednall collection. Accord- 
ing to Davis et ai (1979) there are 3 syntypes \\\ ANSP 
(Shell Cat. No. 67369); we therefore consider the 
selection o( a neotype invalid. Davis et ui (1979) also 
erroneously list the type locality as Cape Yorke 
Peninsula, Qld. 

Since the above specimen came from the Matthews and 
Bednall collection and is from the type locality we 
consider it likely to be a syntype. 

Ischnochiton (Heiemzona) properensis Ashby, 1920 
Trans. R. Soc. S. Aust. 44: 278, pi. 12, fig. 7a-b. 
Holotype: 1)1 1896, entire specimen, from Proper Bay, 
Port Lincoln, S.A., collected by E. Ashby, January, 

Isdinochiton resplendcns Bednall ik Matthews, 1906 

Proc. Malac. Soc. Land, 7(2): 91, pi. 9. figs 4, 4a-f. 

Ischnochiton (Haplopiax) stnaragdinus resplendcns 

Bednall & Matthews, 1906. 

Neotype: D13739, entire specimen with animal, from 

Marino, near Adelaide, S.A., collector and date of 

collection unknown. Selected by Cotton and Godfrey 

(1940: 503, fig. 491). 

Note: Listed and figured by Cotton and Godfrey (1940) 

as neotype without discussion and considered, by us, 

as invalid. 

Ischnochiton (Haploplax) stnaragdinus resplendent 

var. wester nensis Ashby, 1923 

Trans. R. Soc. S. Aust. 41: 226. 

Hololype: DI3722, entire specimen^ from Yallingup, 

W.A., collected by P. Ashby, date a\' collection 


Ischnochiton (Haplopiax) smurugdinus var. tunereus 

Ashby, J 924 

Trans. R. Soc. S. Aust. 48: 3)5. 

Holotype: DII237, enl ire specimen, wilhunimaK from 

Long Reef, N.S.W., collected by W. H. Hatcher, date 

of collection unknown, 

Ischnochiton stroinjelfi Bergenhayn, 1931 
Arkiv. Zool. 23A(I3); II, pi. I, figs 14 16; pi. 2, 
figs 52-56. 

- I.epidochitono stroemfelti (Bergenhayn, 1931). 
Syniype: DI6I86, entire specimen with animal, from 
Luertaventura, Puerto Cabras, Canary Islands, on 
rocks at low ode, collected by N. Odhncr, 1930. 
Note: l-'rorn Ashby collection. 

Ischnochiton ntteunns Bednall, 1897. 

Proc. Malar, Soc Lond 2(4): 147, pi, 12, lig. 3a d. 

Neotype: 1313738, entire specimen with animal, from 

Sultana Bay, Yorke Peninsula, S.A., collector and date 

of collection unknown. Selected by Cotton and 

Godfrey (1940: 493). 

Note: listed by Cotton and Godfrey (1940) without 



discussion, also Davis et at. (1979) list 7 syntypes in 
ANSP (Shell Cat. No. 69143) and we thei^fore consider 
the selection of a ncoiypc invalid. 

hchnochiton tlunvasi Bcdnall, 1S97. 

Pmc. Malac. Soc. Land. 2(4): 149, pi. 12, figs 4a-d, 


- hihnovhiion (Huploplax) thomasi Bcdnall, 1897. 
Neotype: D13737, entire specimen with animal, from 
Manno, near Adelaide, S,A,, collector and date of 
collection unknown. Selected by Cotton and Godfrey 
(1940: 504). 

Note; listed by Cotton and Godfrey (1940) without 
discussion, also Davis et al. (1979) list 8 syntypes in 
ANSP (Shell Cat. No. 69144) and we therefore consider 
the selection of a neoiype invalid. 

Ischnochiton tindalei Ashby, 1924. 
J runs. R. Soc. S. Aust. 48: 323, pi, 31, fig. 4a-c. 
Holotype: D4656, dLsaiticulated specimen with pieces 
of girdle, from Groote Eylandl, Gulf of Carpentaria, 
N.T., on a block o\' dead coral in deep water, collected 
by N. B. Tindalc, date of collection unknown. 
Note: Type unique. 

iepidoplcurus varicgatus Adams & Angas, 1864. 

- Ischttocluton vuricgatus (Adams & Angas, 1864). 

Ischnochiton verconis Ton*, 1911. 
Trans. R. Soc. S, Aust. 35: 102, pi. 24, fig. la-f. 
Holotvpe: DI2868, entire specimen, from Ellcnbrook, 
south of Cape Nat uralistc, W.A., in rockpool on inside 
reef, collected by W. G. Torn Christmas, 1910-11. 
Note: Type unique. 

Genus Ischnonulsia Shuttleworth, 1853 

Ischnorudsiu pupuuensis Ashby, 1923. 

Trans. R. Soc. S. Aust, 47: 227, pi. 17, fig. 2a-c. 

- Isc/wochdon (Ischnoradsju) pupuuensis (Ashby, 

Holotype: D14316, posterior and 2 median valves 
articulated with girdle, resi of specimen disarticulated, 
from Normanby Island, Papua New Guinea, collected 
by R Andrew, date o\' collection unknown. 
Note: Type unique. 

Genus Stenochiton Adams & Angas, 1864 

Stenochiton cymuducealis Ashby, 1918. 

Trans. R. Soc. S. Aust. 42: 70, pi. 13, figs I, 4, 5; pi. 14, 

figs II, 12a-c. 

Holoiype: D98I, entire specimen with animal, from 

Marino, near Adelaide, S.A., on Cymodocea antarctica 

sums, collected by E. Ashbv, date ot collection 


Paratypes: D980, 2 entire specimens with animals, with 
same collection data as holotype, D16545, disarticu- 
lated specimen with posterior valve presumed lost, with 
same collection data as holotype. 
Note: D16545 is labelled "type" but was obviously 
disarticulated for fig. 12. According to Ashby (1918) 
the type is the specimen illustrated in fig. 5 which 
corresponds to the above. 

Stenochiton posidonudis Ashby, 1918. 
Trans. R. Soc. S. Aust. 42: 72, pi. 13, figs 2, 6, pi. 14, 
fig_ I3a-d. 

= Stenochiton pilshryanus (Bcdnall, 1897). 
Holotype: D117I4, entire specimen with animal, from 
Marino, near Adelaide, S.A,, collected by E. Ashby, 
date of collection unknown. 

Paratypes: DU718, 5 entire specimens stuck on one 
card; 2 from Cape Jervis, S.A., collected by E. Ashby 
14.iii.19l8: 1 from Outer Harbour, S.A., and one from 
Marino collected by E. Ashby, date of collection 
unknown and I from Largs Bay, S.A., collected by 
E. H. Matthew;*, date of collection unknown. D16544, 
disarticulated specimen with same collection data as 

Note: DI6544 is labelled "type" but was obviously 
disarticulated for fig. 13. According to Ashby (1918) 
the type is the specimen illustrated in fig. 6 which 
corresponds to the above. 

Stenochiton tatei Ashby, 1927, 
Pap. Proc. R. Soc. Tas. 1926: 113. 
=Stenochifon piisbryanus (Bednall, 1897). 
Holotype: D11729, based on the same specimen as the 
neotypc of Ischnochiton piisbryanus Bednall, 1897, 
selected by Ashby, (1919) and also listed here. 
Note: A new name for the shell described as L 
piisbryanus (Ashby, 1919) as the identification of the 
specimen selected as neoiype was in doubt. 

Genus Tonicel/a Carpenter, 1873 

Chiton mannoreus vai\, caeruteus Winkley, 1894. 
-Tonicella tnormoreus (Eabricus, 1780). 

Genus Trachydermon Carpenter, 1864 

Trachydermon lowei Pilsbry, 1918. 

Nautdus 31(4): 127. 

-Lepidochitona lowei (Pilsbry, 1918). 

Paratype: Dl 1275, entire specimen, from San Pedro, 

California, U.S.A., collected by H. N. Lowe, date of 

collection unknown. 

Note; From Ashby collection. Type in ANSP (Shell 

Cat. No. 117955). 

Trachydermon (Craspedochdus) turtoni Ashby, 1928, 
Proc. Malac. Soc. Land. 18(2): 80, pi. 6, figs _S-8. 


RFC. S. AUST, MUS. IW>: 97-Hi 

June. f&H6 

= Lepidachi(ona Itir toni (Ashby, 1928) 
Holotype: DI0982, 2 median valves, valve fragments 
and fragment of girdle* from (A^rt Alfred, South Africa, 
collected by W. H. Turton, date of collection unknown. 
Note: A label with the valves says "valves not 
photographed 1 *, Rest of type presumed lost, Type 


Genus Lepidopieurus Risso, 1826 

Lepidopieurus badius Hedley & Hull 1909. 
Rec. Aust. Mus. 7: 260, pi. 73, figs 1 & 2. 
^Leptochiton badius (Hedley & Hull, 1909). 
Paratypes: DI0668, 5 entire specimens with animals, 
from Long Reef, near Narraheen, N.S.W., collector and 
date of collection unknown. D12532, one entire 
specimen with animal, with same collection data as 

Note: The localities "Port Stephens'* and "Kangaroo 
Island S.A" have been added to D10668 (in pencil) and 
there is therefore some doubt regarding the type status 
of these specimens, Type in AM (C30459). 

Lepidopieurus colummtrius Hedley & May, 1908. 
Rec Aust. Mus. 7(2): 123, pi. 24. figs 27 & 28. 

leptochiton colutnnarius (Hedley &. May, 1908). 
Paratypes: D10667, 2 median valves, from 7 miles east 
of Cape Pillar, Tas,, dredged with holotype in 100 
fathoms, collected by W. L. May, 18.xii.1907. D15679, 
posterior and 4 median valves, with same collection 
data as D10667. 

Note: D15679 from May collection (No. 226). Type in 
AM (C29060), 

lepidopieurus f inlay i Ashby, 1929. 

Trans, Proc, t\\Z, Inst. 60: 372, pi, 32, figs 5-7. 

- l.epiochiion finlayi (Ashby. 1909). 

Holotype: D11061, 5 disarticulated median valves and 

dried radula, from off Otago Heads, N.Z., dredged in 

60 fathoms by H. J. Finlay, date of collection unknown. 

Paratype: D16I88, entire specimen with animal but 

posterior valve absent, with same collection data as 


Note: Rest of type in AM (€95165). 

Lepidopieurus glauerti Ashby. 1929. 

J, R. Soe. U: Aust. 15: 50, fig. 16. 

= Leptochiton glauerti (Ashby, 1929), 

Paratype: D16152, 5 median valves articulated with 

animal and remains of girdle, anterior valve and other 

median valve disarticulated, from Bathurst Point, 

Rottnest Island, W.A., collected by I.. Glauert, date of 

collection unknown. 

Note: Posterior valve presumed lost, lype in WAM 


Lepidopieurus iredalei Ashby, 1921. 

Proc. R. Soc. J-'/c (NS) 33: 157, pi. 8, fig. 3a-b. 

= Leptocluton inquinatus (Reeve, 1847). 

Holotype. D11240, entire specimen with animal, from 

Doubtless Bay, N.Z., collected by A. F. Brookes, daie 

of collection unknown, 

Lepidopieurus liratus Adams & Angas, 1864. 
Proc. ZooL Soc. Lond. 13: 192. 
= Leptochiton liratus (Adams & Angas, 1864). 
Neotype: D13735, entire specimen with animal, from 
Sultana Bay, Yorke Peninsula, S.A., collected by 
Matthews and Bednall, date of collection unknown. 
Nonet A neotype for L. liratus was selected by Iredale 
and Hull (1925a: 343) and this specimen is in the AM 
(C 10410), The above specimen was presumably selected 
by Cotton and Godfrey (1940: 477, fig. 458) as yet 
another neotype for /„. liratus; however, the specimen 
docs not match fig. 458. The type status of the above 
specimen is therefore extremely doubtful and is invalid 
in any case as Iredale and Hull (1925a) had already 
selected a neotype. 

Lepidopieurus matthewsianus Bednall, 1906. 
Proc Matac Soc Lond. 7: 32, pi. 9, fig. la-f 
^Leptochiton matthewsianus (Bednall, 1906). 
Neotype; D13734, entire specimen .with remains of 
animal, from Marino, near Adelaide, 5.A.* collected 
by E. Ashby, 1919. Selected by Cotton and Godfrey 
(1940: 476, fig. 455). 

Note: Listed and figured by Cotton and Godfrey (1940} 
as neotype without discussion and considered, by us, 
as invalid. 

Lepidopieurus niger Tbrr, 1911. 

Trans. R. Soc, S, Ausr 35; 105, pi. 25, fig. 5a- f. 

= Leptochiton niger (Torr, 1911). 

Holotype: DU686, entire specimen with animal but 

with anterior valve missing, from Hopetoun, W.A., 

under stones in shallow pools, collected by W. G. Torr. 

Christmas, 1910-11. 

Note: Anterior valve presumed lost. Type unique. 

Lepidopieurus pelagicus Torr, ! V >I2 

Trans. R. Soc. S. Aust. 36: 165. pi. 5. fig, 2a-F. 

-Leptochiton columnuris (Hedley & May, 1908). 

Holotype; DII688, anterior, posterior and one median 

valve, dredged in 130 fathoms, off Cape Jaffa, S.A., 

collected by J. C Verco, 25.xii.I905. 

Note: Rest of type presumed lost. 

Lepidopieurus profundus May, 1923. 

Illusi, Index Tas Shells, pi. 14, fig, 2, Appendix 

—Leptochiton profundus (May, 1923). 

Holotype: D12533, entire specimen, dredged in 10 

tcii horns, o(\" Pilot Slat ion, Derwcni River, Tas., 

collected by W. L. May, date of collection unknown. 

Paraiype: D16189, enfire specimen wirh animal, with 



anlericu valve disarticulated, with same collection data 
as holotype. 

Note: May (1923) illustrated L. imfHinatus Syke\ 1896 
but id an appendix says "An examination of the type- 
in the British Museum shows that this is not incjuinatus, 
which is the N.Z. species. Our shell is now to be known 
as profundus Ashby'\ Unfortunately Ashby's 
description of profundus was published after May's 
and in any case is a different species. 

I, epidopleurus profundus Ashby, 1923. 

Trans, R, Soc. S, Aust. 47: 221, pi. 16, figs 2, 2a. 

= Leptochiton co/lusor (Iredale & Hull, 1925). 

Holotype; Dl 1288, entire specimen with animal, from 

Gulf St Vincent, S.A., dredged by .1. C Verco, date of 

collection unknown, 

Paratopes: D1I691, 2 specimens, one entire, the other 

with animal but with anterior, posterior and one 

median valve disarticulated, from Port Phillip Bay, Vic., 

dredged by Bracebridge Wilson (No. 881), date of 

collection unknown. 

Note: Renamed Parachiton cottusor by Iredale and 

Hull (1925a), also listed here, as L. profundus was 


l.epidopleurus variegafus Adams & Angas, 1864. 

Proc. Zool. Soc. Land. 13: 192. 

= Ischnochiton variegafus (Adams & Angas, 1864) 


Neotype: D13736, entire specimen with animal, from 

Minlacowjc, Hardwicke Bay, Yorke Peninsula, S.A., 

collected by E. H. Matthews, date of collection 

unknown. Selected by Iredale and Hull (1927: 13). 

Genus Parachiton Thiele, 1909 

Parachiton collusor Iredale & Hull, 1925. 
Aust. Zool 3(8): 346, pi. 39, fig, 22. 
^Leptochiton collusor (Iredale & Hull, 1925), 
Holotype: D11288, based on the same specimen as the 
holotype of Lepidopleunts profundus Ashby, 1923, also 
listed here. 

Note: There is some doubt that this specimen was the 
one selected as type by Iredale and Hull (1925a) as 
Iredale and Hull (1927) list the type of P. cullusor in 
SAM and the type of L, profundus in the Ashby 
collection. However, later labels with the specimen say 
that it is the type of R collusor and Cotton and 
Godfrey (1940) also list and figure it as the type. 

Parachiton venvnis Cotton & Weeding, 1939, 
Trans. R. Soc. S. Aust. 63(2): 183, pi. 7, fig. 2. 
-Leptochiton verconis (Cotton & Weeding, 1939). 
Holotype: D11689, posterior valve only, from near St 
Francis Island, Nuyts Archipelago, S.A.. dredged in 
15-20 fathoms by J. C. Verco, date of collection 
Note: Type description was based on this single valve. 

Genus Terenochiton Iredale, 1914 

Terenochiion iscus Cotton & Weeding, 1939- 
Trans. R. Soc. S. Aust. 63(2): 182, pi. 7, fig. 1. 
= Leptochiton iscus (Cotton & Weeding, 1939). 
Holotype: D1232, entire specimen with animal, from 
Cape Jervjs, S.A., collected by F L. Saunders, 1917. 


Genus Kopionella Ashby, 1919 

Kopionella matthewsi van intermedia Ashby, 1927, 
Pap. Proc, R. Soc, Tas, 1926: 10]. 
- Plaxiphora (Fremblya) maithewsi (Iredale, 1910). 
Syntypes: 0121 1 5, 3 entire specimens with animal 
remains, from Penguin, north-western Tas., collected 
by E. Ashby, November, 1924. 

Kopionella iasmanica Ashby, 1920. 
Trans. R, Soc S. Aust, 44: 268, pi, 11, fig. la-d. 
-Plaxiphora (Fremblya) matthewsi (Iredale, 1910). 
Holotype: D12156, disarticulated specimen, from 
Lunawanna, South Bruny Island, D'Fntrccasteaux 
Channel, Tas., collected by E. Ashby, March, 1920. 
Note: Smith and Robertson (1970) list the holotype as 
lost as it couid not be located at the time. 

Genus Plaxiphora Gray, 1847 

Plaxiphora hedieyi Torr, 1911. 

Trans, R, Soc S, Aust, 35; 103, pi. 24, fig. 2a-f, 

-Plaxiphora (Fremblya) maithewsi (Iredale, 1910). 

Holotype: D1287I, entire specimen with anterior and 

posterior valve disarticulated, from Rabbit Island, near 

Albany, W.A., collected by W. G. Torr, Christmas, 


Plaxiphora pustulosa Torr, 1911. 

thins* R. Soc S. Aust 35; 107, pi, 25, fig. 7. 

- Uolophura (CtovarizonaJ hirtosa (Blainville, 1825) 


Holotype: DI3719, one median valve only, from 

Albany, W.A., collected by W. G. Torr, Christmas, 


Note: Type description was based on this single valve. 

Plaxiphora zebra Torr, 1911 

Trans. R. Soc, S. Aust. 35: 106, pi. 25, fig. 6. 

= Plaxiphora (Fremblya) matfhewsi (Iredale, 1910). 

Holotype: D12869, one median valve only, from Port 

Esperance, W.A., collected by W. G. Torr, Christmas, 


Note: Type description was based on this single valve. 


RET. V Al ST Mbs\ lt?(«P 07 115 

.////><•, /y,v6 


Genus Lorkella Pilsbrv, IS93 

Genus Lariat H & A Adams, 1852 

Loriea e/liottae Coiton & Weeding, 1939 
Tram. R. Soc. S. Aust. 63(2): 189, pi. 7, fig. 9. 
HoloLype: D11658, entire specimen, from Rottnest 
Island, W.A., collected by L. A. Elliott, January, 1933. 
Note; Type unique. 

l.orieella torn Ashby, 1919 

Trans. & Soc. S Aiis/. 43: 62, pi. 10, Fig. 16. 

- !.oncella QffgOSi (H, Adams in H. Adams A Angus, 


Holotype: D12440, entire specimen, I'rom Quaranline 

Station, Port Jackson, N.S.W., in shallow water ai low 

Tide, collected by E. Ashby, November, 1918. 

Loriea haurakiensis Meslayer, 1921. 

Trans. Pmc. VZ. Inst. 53: 177, pi. }^ % figs 1-3. 

Paratype: D1I089, entire specimen, dredged in 20 

fathoms, olT Kauwau and Tiritiri islands. Hauraki 

Gulf, near Auckland, N.Z., collected by A. E Brookes, 

date of collection unknown. 

Note; Type in NMNZ (M1121). 


Genus Subterenochiton Iredalc & Hull, 1924 

hvhnochiron bednalli To it, 1912. 

= Subterenochiton bednalli (Tort, 1912). 



\SHHV, F. 1918. Monograph of llic yeruis Sfc/mciiituti (Order 

Pohnlacophora). Willi descriptions of I wo new >pccies, 7'nttts. 

A\ Soc. S, Aust. 42; (o-7N, pis 13 & 14. 
ASHBY, fci 1919. Notes on Australian PolypUiujphora. including 

descriptions of two new genera, a new wuietv. and (lie 

description and proposed recognition ol Mi. Bidnatl\ 

SU'OOchltint pifshryanas, Trans, R. Sue, .S. Aust 43: 66-73, pi. II. 
ASHIJY, P.. 1922, Tvpes o\ .species of Australian Polyplaeophora 

descrihed by De'Blaiinille, I amarck, Dc'Roclibiuni:, and others, 

now in the Museum d'Histoire Naturellc, in Parts. Trans, ft Soc. 

S. Aust 4ft: 572-5H2. 
ASHBY. L, 1926. The Aeanihoid Chalons ol New Zealand, with 

descriptions and figures, including several new specie*, flvti 

Value. Soc. Loud. 171: 5-34, pts 1-4. 
ASHBY. \ IQ27, Notes on, .\\m\ additions lo, the Chiton fauna of 

North West Tasmania, together with a brief review of lite genus 

Srcuochittnt Hop. Hroe A*. Soe Tasnt 1926: 92-117. 
ASHBV, L. 1928. Notes on a collection ol' chitons (Polyplaeophora) 

Irom the Capricorn Group. Queensland. Truth. /?, Sue. S Au\i 

52: 167 173. pi. 12; 
ASI IRY, E. 1931. Monograph of I lie South African Polyplaeophora 

(Chitons). Amt S. Aft \ticv MHU: J-5S>, pis. 1-7. 
ASIIUY. E„ and COTTON, B. C. 1937. Descripiion of two new 

\pecK-s ol Austiiihan Chitons with additional notes and leeords. 

train, H Soc, V Aust. 41: 145-148. pi- X 
UFRC.FNHAYN. I. R. M 19.51. Beiiragc /.in Malako/oologic der 

Kanarisehen Inseln. Die Loricaten Arkiv. Zoo/. 23A(I3): I *S; 

pis. 1-3. 
COl ION, I?. C. 1954, South Australian Molltrjca: Chi/otts: Govt. 

Printer Adelaide, 151 pp, 
COTTON. B. C andOOUl -RLY.KK. W40. The Molluscs oj 'South 

Australia fart //, Scaphi)poda, Ct/>lutlopt.*da, Aplacopohora 

atnt C'repipodu. Oovt Prime/ Adelaide, 2<N4 pp. 

COTTON. B. C N and WLLDING, B. J, 1939. I fodtttUn I oricalcv 

ham ft Soc. S. Aust 03: 180-199, pi. 7 
OAYls. G. M.. KOBLRrSON, ft., and Mil LER, M. W§, I aialofiic 

al iIr 1 Cniloti types nf rhe Acadfirfly ot Natural Sctonccs ot 

Pliiladelphi.i. Trvo/uu I l-ft0_ 
DL PLdS, p l'.'17. Noics piiscsMU Luiirsde IVxanicn de la Collection 

dc Polvplaciiplires clu Museiin) de Paris. Hull. Mus. \'utn. ffht 

nut tons 2.1: 553 H 
l)t. PLilS. P. 1 918. Notes coneernanT les Polvplacophores. Hull Mus. 

Slot. Mist Slut. 24: 525 533. 
QU LS. L., and OOSI.INFR. I, 19S3. Primary type specimens o\ 

marine mollusca (e\Llnding c ephalopod.-n in the Somh African 

Museum. Aon V. \fr Mus. 92: T-.S2. 
IRLDAPL, T, and Hill I, A. I'. B. 1925a. A Vlonorraph ..' 

Australian loricates, pari IV. Aust. loot. 3: 339-362. pis 

IRl-.n.AIF, 1., and lltJLP, A. I-. B. I925K A ot 

Australian Lorieatc\ Pari V, Aust /oof 4: 75 I H. pis 9-12. 
IRLDAIl:. T.. and HUM, A. I 11 1V27. ,| Moiioy/uph of 

Australian loricate f/-7n turn Mollusca Order i. oriental. R 

/ooL SitL. N.SAV. publ.. 168 pp., 2) pjftlttS 
KAAE, P. and VAN BLI-LP, R. A. 1980. Cotaiuytw of Living 

Chitons. I")r \V. Backhuvn Pul>l., Rotictdam, 144 pp. 
M^tPIIAlL, M. K.. and /Hl)l FR. V\. 197ft. H.stoi) in u 

Tiismanian Chiton. Jits. Afirf, No, 53: I", 
MAUCjIIAN, M. M. 19t)0. UeriiHiion ot a n»W species of South 

Australian Polyplaeophora. Trans. A 1 Snv. S' Aust 24:89, pi, I 
MAY, W. L. 1923. '1// fliustratctl hutcX t>J Tasioanuio Shells. CjOVt- 

Printer Hoharl. 54 pp., 50 pis., Appendix. 
SMITH. H. J, and ROBLRTSUN, K C. I97(). C. atalojjut of( hiion 

(Vitiptiiiiciiai, \tiillusL;i) lypc^ lit (lie National Muvnim ill 

\it.ioria, Australia. Meat Nat. Mas. \ ie. 31: Kl 9tt, 


List of Chiton "types" from the Dupuis collection 
given to Ashby by Dupuis. All axe labelled "TYPE" 
or "CO-TYPE" usually with a Dupuis label with the 
word "TYPE" in red print, glued to the label and the 
prefix "eu" added by hand. The specimens are listed 
in alphabetical order according to the original name 
at the time of description. 

Acanthopleura balansae Rochebrune. 1882. 
Bull. Sac. Phitom. Paris, Ser. 7, 6: 197. 

Ac(7/nho/)iettra liadtiuni Wink worth, 1927 

Syntype: D10231, entire specimen with animal, from 
Timor, collector and date of collection unknown. 

Note: Specimen with label "co-TYPE de A. balansae 
Roch! f and reidentilied as /I. spim'gera Sowcrby. "the 
locality given by Rochcburne (1882) is "Australie (Peron 
et Lesueur). Nouvelle Caledonie (Halansa; Germain)" 
The type status of the above specimen is therefore very 

Chiton elongatus Blainville, 1825. 
Diet. Set. Nat. 36; 352, 

fsJitiochiton etonxatas (Blainville, 1825) 

Syntype: D10237, entire specimen, from King Island, 
Bass StraiL collected by Peron and Lesueur, 1802. 
Note: The shell is marked on the inside "tie King 11 



presumably in the handwriting of Peron or LcstieiiT and 

is a historically significant shell (Maephail & Zeidler, 
1978). Jt was sent to Ashby by Dupuis as "co-types" 
of/, iineolatus (Ashby, 1922) and a Dupuis label says 
the same. Ashby (1922) determined it as /. crispus 
(Reeve, 1847) which is now considered a synonym ot 
/, etongatus, The above specimen matches ihe original 
desci iption of C. elongatus and considering that it was 
collected by Peron and Lesueur it was most likely pari 
of the series of specimens originally seen by BlainvilJc. 
Syntypcs: D10238, 3 entire specimens with same 
collection data as DI0237. 

Note: = /. suhviridus (Iredale & May, 1916) but would 
have been considered f, elongatus in the past. There 
is no original label with the specimen but a label 
presumably Written by Ashby says "col. by Peron and 
Lesueur> Is, King 1802 (with Biainville's type of 
lineola(usy\ If the information with the specimens is 
correct then they could also have been part of (he 
original material seen by Blainville. 

Chiton iumvt Dupuis, 1917. 
Bull. Mus. Natn. Hist, flat Paris 23: 538. 
-Chiton peregrinus I hielc, 1910 (CHITONIDAE). 
Syntypcs: D102U, 10 entire specimens, only one with 
animal, from Aden, Red Sea. collected by Dr 
Jousseume, date of collection unknown. DI0252. one 
disarticulated specimen with piece of dried girdle, from 
the Red Sea, collected by Dr Jousseume, date of 
collection unknown. D10255, one disarticulated 
specimen with piece of dried girdle with same collection 
data as D10252. D11200, one entire specimen from the 
Red Sea, collector and date o\ collection unknown. 
Note; All of the above are labelled "TYPE" except for 
D1 1200 which is labelled "co-type". As Dupuis did not 
clearly designate a type and since the above specimens 
were given to Ashby by him, il is likely that they are 
all syntypcs. 

Chiton tehuelchus D'Orbigny, 1841. 
Voy Amer. merit!., Moll. 3(3); 488, pi. 65, figs 7-13. 
- Chad o pleura an gu lata (Spengler, 1797) 

Syntype; DI0293, entire specimen, from Bay of San 
Bias, Patagonia, collector and date ot collection 


Note: The above specimen is from the type locality but 
in the absence of original labels the type status must 
be very doubtful. 

Chiton undulatus Quoy & Gaimard, 1835.. 

Voy. de (Astrolabe, Zool. 3: 393, pi. 75, figs 19-24. 

= Oni/hochiton neglect us Roehebrune, 1881 


Syntypcs: DI0223, 2 entire specimens, from N.Z., 

collector and date ftf collection unknown. 

Note: Specimens of very doubtful type status. The type 

is in MNHN and was seen by Ashby (1922). However, 

Quoy and Gaimard recorded several specimens but 
apparently did not designate a type so the atftJVC 
specimens could have been pari of Ihe type scries, 

Chiton violaceus Quoy & Gaimard, 1835. 

Voy. de ('Astrolabe, Zool. 3: 403, pi, 73, figs 15-20, 

-Notopkix violaceus (Quoy & Gaimard, 1835) 


Svntvpe: D11054, disarticulated specimen, from NX, 

collector and date of collection unknown. 

Note: A specimen of doubtful type status. The Dupuis 

label with the specimen has "Tasman Bay nr. Nelson" 

added in pencil, which is the type locality. 

Cryptoptax calcdonicus Roehebrune, 1882. 

Bull. Soc. Philom. Paris, Scr. 7, 6: 196. 

= Cryptop(ax larvaefonuis (Burrows, 18)5) 


Paratypcs: D10330, 3 entire specimens with animals, 

from New Caledonia, collector and date of collection 


Note; A Dupuis label wiih the specimens designates 

them as "co-types" but a later Ashby collection label 

has the words ''with TYPE 11 on it thus indicating that 

the above specimens eouW be paratypes. 

Cryptoconchus stewartittnus Roehebrune, 1882. 

ML Soc Phdom. Pans, Ser. 7, 6; 194. 

= Cryptoconchus ])omsus (Burrows, 1815) 


Para type: Dl 1053, disarticulated specimen, from Ciurk 

Strait, N.Z., collector and date of collection unknown. 

Note: The above specimen is most likely a paratypc as 

Ashby (1926) described and illustrated it and referred 

to it as "one of Rochcbrune's colypes". The type, a 

spirit specimen, is in MNHN (Ashby 1922). 

Gymnoplax spicijerus Roehebrune, ISM. 
Bali Soc Philom. Paris, Ser. 7, 8: 36. 

Ischnochiton (Ischnoplax) pectinatus (Sowerby, 
Syntype: D10297, I ! /: median valves, from Cochmo 
Island, Guadeloupe, collector and date of collection 

Note: The above material is from the type locality and 
was probably part of the type series. 

Lepidoplearus eatnphelii l-ilhol, 1880. 

Comptes Rendus hebd. Seanc. Acad. Sci. Paris, 91: 


= Ischnochiton circumvailatus (Reeve, 1847) 


Syntype: DU022, animal and disarticulated valves, 

from Campbell Island, N.Z., collector and date of 

collection unknown. 

Note: There is some doubt that this is one o\' Filhol's 

types; however* it matches the description given by 

Dupuis (1917) for specimen "B" oi' two specimens that 


Kle. S. AUSI. MUS. 19(8): 97-115 

June. 1986 

he regarded as type material of L, camphelli. There is 
also a radula slide With DU022 but ii docs not belong 
to the above specimen; however, the SAM register refers 
to two specimens one of which may have been used 
for the radula preparation. 

Lepidopleurus cessaci Rochebrune, 1881. 
Ball. Soc. Philom, Paris, Ser. 7, 5: 118. 
-Ischnochiion cessaci (Rochebrune, 1881) (ISCHNO- 

Syntype: D10324, disarticulated specimen, from Cape 
Verde Island, collector and date of collection unknown. 
Note: SAM records indicate that there were another 
S specimens which Ashby kept and were presumably 
lost in the tire at Ashby's house. The above specimen 
is from the type locality and was probably part of the 
type series. 

lepidopleurus fodiatus Rochebrune, 1881. 
Bull. Soc. Philom. Paris, Ser. 7, 5: 119. 

- Ischnochiion textilis (Gray, I&281 

Syntype: D10286, entire specimen, from Natal, South 
Africa, collected by M. Verreaux, date of collection 

Note: The type status of this specimen is in doubt. The 
label with the specimen reads "Ischnochiion tigrinus 
Krauss (eo-TYPE dc Lepidopleurus fodiatus Rocheb. 
Australia!!) Verreaux NatalV\ Obviously there is some 
confusion regarding the locality data. 

Ashby (1922) says that he has never seen this species 
in Australia and is confident thai New Holland, the 
type locality, is erroneous. 

Lepidopleurus melanterus Rochebrune, 1884. 
Bull. Soc. Philom Paris, Ser. 7, 8. 37. 

- Ischnochiion circumvallatus (Reeve, 1847) 

Syntypes: 1)10292, 7 entire specimens with animals, 
from Campbell Island, N.Z., collector and date of 
collection unknown. DI03J9, one entire specimen with 
animal, with same collection data as D10292. D13061, 
one entire specimen with animal, with same collection 
data as D10292 

Note: The above material is from the type locality and 
at least DI0292 could have been part of the type series. 
D103I9 and D13061 are without the usual Dupuis label 
and may have been extracted from D10292. 

Lepidopleurus rochebruni Jousseaume, 1893 

Bull. Soc. Philom. Paris, Ser. 8, 6: 102. 

= Calltstochiron adenensis (E. A. Smith, 1891) 


Syntype: DI0326, one entire specimen, from Djibouti, 

Gulf of Aden, collector and date of collection 

unknown. DI0342, 2 disarticulated specimens, one 

from Djibouti, the other from Aden, Gulf of Aden. 

collector and date of collection unknown. 

Note: DJ0326 is clearly marked "cotype du Dr 
JtftfSsesafJrai^ and the specimen matches the description 
and the largest measurement given by Jousseaume. It 
is therefore most probably a syntype. The Type status 
Of DI0342 is rather more doubtful. 

Noiochiton mirundus Thiele, 1906 
Hiss. lirgebn. d. Tie/see Exp 9(2): 332, pi. 29, figs 

=-Nuttalloehtton mirundus (Thiele, 1906) (ISCH- 

Syntype: D102I3, 5 median valves articulated with 
girdle and disarticulated anterior, posterior and median 
valve 2, from Antarctica, collector and dale of 
collection unknown. 

Note: Despite the fact that D10213 is clearly labelled 
"cotype" it cannot be Thiele > s type of N. mirandus&s 
Thiele mentions only one specimen and an anterior 
valve from cast of Bouvet Island (Valdivia Stn. 127) 
and this specimen was several times larger than D10213. 

Onithochiton filholi Rochebrune, 1881 

Bull. Soc. Philom. Paris, Ser. 7, 5: 120. 

-Onithochiton neglect us Rochebrune, 1881 


Syntype: D10215, disarticulated specimen, from Cook 

Strait, N.Z., eolleetor and date of collection unknown. 

Note: Clearly labelled "co-TYPE de O. filholi Roch". 

Onithochiton neglectus Rochebrune, 1881 


Bull, Soc Philom, Paris, Ser. 7, 5: 120. 

Syntypes: D10221, two entire specimens with animals, 

without collection data. 

Note: In the absence of original labels and good 

locality data it is difficult to ascertain the type slants 

of the above specimens. 

Schizochifon hyadesi Rochebrune, 18S9 
Miss. Sci. Cap. Horn 4 ZooL, : 132, pi. 9, fig. 1. 
= Nultallochilon hyadesi (Rocheburne, 1889) (ISCH- 

Syntype: D10189, entire specimen with animal, from 
TeiYa del Fuego, collector and date of collection 

Note: The type status of this specimen is very doubtful. 
The original label does not indicate the type status but 
the words "cotype-paratype* 1 have been added in pencil 
on the back of the label. 

Schizochiton jousseaumei Dupuis, 1917 (SCHIZO- 

Ball, Mus. Main. Hist. natr. Paris, 23. 536, figs t-4. 
Syntypes: D10233, 2 entire specimens with animals plus 
one anterior valve, from the Red Sea, collected by 
Jousseaume, date of colleetion unknown. D10235, one 
entire specimen with animal with same collection data 
as D 10233 



Note: Although the above specimens are labelled 
"TYPE" it is unlikely that any of them are types as 
the species was based on a unique specimen which is 
in MNHN. The single anterior valve of D10233 is 
labelled "Perim", the type locality. 

Tonica fontainei Rochebrune, 1882 

Bull. Soc. Philom. Paris, Ser. 7, 6: 193. 

= Ischnochiton punctulatissimus (Sowerby, 1832) 


Syntypes: D10291, 4 entire specimens with animals, 2 

disarticulated specimens plus 3 end valves and 4 median 

valves from at least 2 specimens, from Chile, collector 

and date of collection unknown. 

Note: The specimens are clearly marked "co-TYPE" 

and are probably types. 

Tonicia lebruni Rochebrune, 1884 
Bull. Soc. Philom. Paris, Ser. 7, 8: 35. 

Syntype: D10196, entire specimen with animal, from 
Patagonia, collector and date of collection unknown. 
Note: The specimen is labelled "TYPE" and in the 
absence of evidence to the contrary it must be 
considered possible type material. 


We wish to thank the following for supplying us with 
information regarding types held by their respective 
institutions: Dr W. O. Cernohorsky, AIM; Mr I. Loch, 
AM; Dr B. A. Marshall, NMNZ; Mr J. Stanisic, QM; 
Dr A. M. Testud, MNHN and Dr F. E. Wells, WAM. 

We are especially grateful to Dr M. K. Macphail who 
initially curated the mollusc type collection under a 
State Unemployment Relief Scheme. We also wish to 
acknowledge Mrs Jan Forrest, SAM, who took the 
photographs and prepared Figure 1 and Miss J. Connor 
who typed the manuscript. 



byW. ZeidlerandF. W. E. Rowe 


The status of the genus Nectria is discussed and the genus transferred from the family Goniasteridae 
to the family Oreasteridae. A previously closely allied genus, Nectriaster, is also transferred to the 
Oreasteridae. The status of the subfamily Nectriinae is discussed and maintained for the unique 
genus Nectria. A new subfamily, Oreasterinae, is erected for the remaining genera of the family 
Oreasteridae including Nectriaster. Problems relating to Gray's (1840, 1847a, b) misunderstanding 
of N. ocellifera which he based on Oudart's (1827) figure of Lamarck's species are discussed and 
Oudart's figure is reproduced for the first time. Eight species are currently recognised including a 
new species from Tasmania. Each of the eight species is diagnosed and geographic ranges detailed. 
A lectotype is selected for N. ocellata Perrier. N. pedicelligera Mortensen is recognised as a valid 
species restricted to the southern Australian coast. The species potential of the genus is considered 
not yet to have been reached. Nectria is recognised as an endemic southern Australian genus. 



W ZEIDtER 1 and F. W. E. ROWE* 

South Australian Museum, North terrace, Adelaide, South Australia 5000 
Die Australian Museum, 6-8 College Street, Sydney, New South Wales 2000 

(Manuscript accepted 5 Murch 1985} 


ZLIDLl-K, Wm ami feCftVE, f . VV E. 1986. A revision of the southern 
Ausualian siarl'ish genus \'txtrui (AiteroUleti: OnMsieinJaej, 
Aiih the description of J new species. /?«v. 5. Aust. Mus. I'J(9>: 
The status of the genus Nectria is discussed and the 
genus transferred from the family Goniasteridae to tlie 
family Oreasteridae. A previously eloscly allied genus, 
Neetriaster, is also transferred to the Oreasteridae. The 
status of the subfamily Nectriinac is discussed and 
maintained for I he unique genus Nectria, A new 
subfamily, Oreastcrinac, is erected for the remaining 
genera of the family Oreasteridae including Neetriaster, 
Problems relating to Gray's (1840, 1847a, b) misunder- 
standing of fa ocellifera which he based on Oudart's 
(1827) figure of Lamarck's species are discussed and 
Oudarl's figure is reproduced for the first time. Eight 
species arc currently recognised including a new species 
from Tasmania. Each of the eight species is diagnosed 
and geographic ranges detailed. A lectotype is selected 
for V. oceliatu Petricr. N. pedicel libera Mortcnsen is 
recognised as a valid species restricted to the southern 
Australian coast. The species potential of the genus is 
considered not yet to have been reached, Nectria is 
recognised as an endemic southern Australian genus. 


Although Nectria is a well established southern 
Australian genus and is easily recognisable, Lhcre has 
been some contusion regarding the type species and 
the identity of a number of the species despite ihc 
reviews of Clark (1966) and Shepherd (1967), 

We have examined all available type material for each 
of the species together with the collections held in the 
Australian Museum Sydney (AM); Museum of 
Victoria, Melbourne (MV) (specimens not measured); 
South Australian Museum, Adelaide (SAM); Western 
Australian Museum, Perth (WAM); Tasmanian 
Museum and Art Gallery, Hobart (TM); Queen 
Victoria Museum and Art Gallery, Launceston (QVM) 
(specimens noi measured) and Museum of 
Comparative /oology. Harvard. U.S.A. (MCZ). 
Following our examination of this material it is 
apparent that Clark (1966) and Shepherd (1967) did not 
appreciate the limits of the species. 

We have also obtained a copy of Oudarr\ (1827) 
figure oi'Asterias ocelli/era Lamarck 1816 (there is no 

accompanying text) which was used by Gray to identify 
his specimens when he established the genus Nectria. 
This figure has apparently noi been sighted since 
Pettier (1875) and we reproduce it here (Fig. I) in order 
to clarify the confused situation created by Gray's 
(1840, 1847a, b) use of this figure. 

We arc now able to recognise eight species of Nectria 
one of whieh is new to science. We are further able to 
detail the distribution of these eight species. 

Nectria is transferred to the family Oreasteridae but 
the status of the subfamily Neetriinae is maintained. 

The following additional abbreviations are used in 
the text. 

BMNH -British Museum (Natural History) 
MNHN = Museum National D'Histoire Natureile, Pari*. 
NSW =New South Wales 
SA -South Australia 
Tas =Tasmania 
Vie = Victoria 
WA ^Western Australia 


Family OREASTERIDAE Fisher, 1911 

Subfamily Neclriinae Pertier, 1894 

Genus Nectria Gray, 1840 

Nectria Gray, 1840(1841): 287; 1866: 15; Dujardin 
& Hupc, 1862: 406; Perrier, 1875: 185; 1876: 1; 1894: 
333; Viguier, 1878: 187; H. L Clark, 1946: 85; A. M. 

Clark, 1966: 309, 

Diagnosis: An oreasterid genus with 5 tapering arms 
(R up to 125 mm) and relatively large disc (r up to 
45 mm). Abactinal surface convex, aetinal surface Hat. 
Abactinai skeleton comprises distinctive raised tabulae 
each supported by 6 radiating ossicles at the base 
forming a network. Disc tabulae crowned with 
peripheral ring of usually prominant granules encircling 
central granules, remainder with granules Of varying 
shape and size. Tabulae distinguished on disc, 
decreasing in si/e along arm to about \fa K after which 
they become unrecognisable as tabulae. Marginal plates 
obvious, Hat, squarish, infero- and supero-marginals 
of similar size and number decreasing regularly in size 


REC. S. AUST. MUS. I9(fte 117-138 

Al#4 BWfi 

lo arm tip, covered with evcn-si/ed granules. Actinal 
plates up to 6-8 rows at base of rays, decreasing di.stally, 
covered with coarse granules. AdambulacraJ plates with 
furrow spines being Mule larger than adjacent granules 
on actinal plates. Papulae in discrete groups between 
radiating ossicles of dorsal skeleton sometimes 
occurring between marginal plates but rarely on actinal 
surface immediately below inferomarginal plates, 
[nterradial septae partially calcified. Pediceliariae often 
present (Fig. 2). Superambulaeral plates present. 

Nectria is an endemic Australian genus occurring 
from the mid-west coast of Western Australia 
(Houtman Abrolhos) along the southern coast to 
Uroughton I. off Port Stephens, N.S.W. in the east (Fig. 
3). Records of specimens from Fiji (Sladen 1889), New 
Zealand (Moriensen I925> and Mauritius (MV 
specimen) are erroneous. Species oceur from the 
shoreline down lo 550 m and are recorded from habitats 
associated wilh rocky reels. 

Remarks: The genus was established by Gray (1840) 
based on a specimen or specimens without locality data 
which he identified as Asterias ocellifera Lamarck, 1816 
(as "aculifera" lapsus cal.) using Oudart'.s (1827) figure 
(Fig I) of Lamarck's species. However we have 
determined that of the few dried specimens of Nectria 
in the BMNH from the mid-!9th century likely to have 
been seen by Gray (1840), none are N ocellifera. One 
specimen in particular (BMNH; 1953; 4.27.24) (Fig. 6b) 
which was certainly seen by Gray fc *as its oldest label 
has 'Nectria ocalifera* stuck on the back of the board 
with l = Asterias ocellifera Lamarck* below and 'Gray' 
pencilled after ocutifera" (A. M. Clark, pers. comm.) 
is cleariy N. ocellata 

Thus like Perrier (1875) we conclude that Gray ( IS40) 
mistook one species for the other. Clearly Gray never 
saw a specimen of N, ocellifera for later (1847a, b) he 
identified a specimen as Patiria ocellifera, noting that 
he may have mis-identified the BMNH Nectria since 
he says M lhis species [P ocellifera] more nearly 
resembles Oudarfs figure than the species 1 have 
described under the name Nectria oculifera y \ Jn shape, 
but not structure, N. ocellifera resembles P. ocellifera 
more so than does N ocellata. We regard Gray's (1847a} 
Patiria ocellifera another mis-identification based on 
Oudart's figure and not a description of a new species, 
However, Clark (1963) has redescribed and figured 
Gray's specimen of P ocellifera as the holotype of that 
species since she does not consider Gray mis-idcntificd 
the specimen 

The above confusion raises the problem of whether 
ocellifera or ocellata should now be the type species 
of the genus Although H. L. Clark (1946) has stated 
'Asterias ocellifera is the type species of Nectria wc 
concur with A. M. Clark (1966) in that "strictly 
speaking, N. ocellata Pcrrier might be considered as 
the type species since that is the one that Gray had 
before him when he diagnosed the genus Nectria, 
though he mistakenly identified ii with Lamarck's 

species* 1 According to Article 70a of the International 
Code of Zoological Nomenclature (ICZN) a decision 
on such a matter should be left to the Commission. 

Nectria was first placed in the family Pentaecrotidae 
by Gray (1840); subsequently Perrier ($75) included 
the genus in the Goniasteridae, dropping all reference 
to the Peiitaeeroiidac*. Later (1894) Perrier placed the 
genus in Its own subfamily the Ncctriinae on ihc basis 
of the distinctive tabulae. Following the e\aminaiion 
of two specimens Fisher (1911) maintained the sub- 
familial status in the family Goniasteridae on the basis 
of the presence of inlet marginal papulae which he 
considered a most important character; he also 
described the occurrences of superambulaeral plates m 
a number of goniasterid genera including Nectria. 
Spencer and Wright (1966) uphold Necrriinae On the 
same basis as Fisher (1911). They include wirhin the 
subfamily one other genus, Nectriaster H L. Clark, 
1946. However, we transfer both of fhese genera to the 
tamiiy Oreastcridae, since, like such genera as Oreaster 
and Penracerasrer, Nectria has a substantially complete 
calcareous intcrbrachial septum, and Nectriaster 
possesses a complete, calcareous intcrbrachial septum 
The inierbrachial septum of goniasterids is rypieally 
membranous. Also, in both Nectria and Nectriaster, 
the alignment of marginal plates, their granulose 
covering, and the arrangement of abaetinal plates in 
particular, are similar 10 other oreasterids. 

The relationship between Nectria and Nectriaster 
does not appear as close as that suggested by H. L. 
Clark (1946) and subsequent authors. Indeed the dorsal 
armament of Nectriaster is not unlike that of Oreaster 
and Petitaeeraster, to which it may be considered more 
closely aligned. However, the distinctive abaetinal 
tabulae of Nectria stands it apan from all other 
oreasterids and for this reason wc prefer to retain the 
subfamily Nectriinae, as did Perrier, but within the 
family Oreastcridae. With the recognition of this 
subfamily foi the type-genus Nectria, the remaining 
genera within the Oreastcridae should be referred (until 
otherwise revised) to the nominative subfamily 
Oreasierinae subiaimnov. which is diagnosed herein 
as a subfamily of Oreasteridae which lacks tabulae and 
for which Oreaster is nominated as the type -genus. 

Clearly, however, the family Oreastcridae is in need 
of critical reappraisal. 


1 Inremuirginal papulae present, when absent many distal 

arm plates enlarged 2 

— Jutcrmurpiiial papulae absent, distal aim plates ncscr 

enlarged . . . . , ,......, 5 

2 Tahutac with concave aspect, with central granules lower 

in profile and smaller than pcriphab 

- ------ . - . V. tmicrubnichid 

— Tabulae with comes, aspect, with centra! granules higher i\\ 

profile and larger than peripherals 3 

to VISION (>l UILU-NI'S \hcrfi/.\ 


3 Dorsal arm plates depths tegularly in sue towards arm angular jn crosvsection. Peripheral granules radiating, 

tip; pediecllariae- with slender valves .*";••• about twice as long as wide, up to I mm high, thin, 

; * - N - pertteWft*w scale-like, taperihfi to rounded tip. Tabulae more or less 

almost as lame as disc plales; pediecllariae with slender .... 

or broad valves 4 decreasing in si/e towards arm tip, with low granules 

4 furrow spita 4-6 (rarely .1 or 7); pediecllariae common giving smooth covering, peripheral granules reduced 

wilh broad valves; huei marginal papulae always present mn distinctive, centrals larger than peripherals', limits 

v nnif/ ^P ,iia [ plates distinct to naked eve. Marginal plates distinct, 

-Furruw spines J-4 (rarely 2) Mi C cllaria,n.,v ^.h^endor s • ^ bu( ^ 

valves; mtermargmal papulae sometimes absent ... , , 

N. utoria compact, distinctly smaller than those on uctinal or 

5 'tabulae low annul I mm high ... H huniilis sp.nov. abactinal surfaces. Actinal plates up to 6 rows 

— Tabulae well developed more lhan 1 mm high .... 6 proximally. Furrow spines 2-4. Subambulacral spines 

6 Marginal plales whh granules disiiucily smaller than on 34 on ^ Rnv apd j,j on second r(m p apu [ ilc 

aetinal surtaee; tabulae with very low, Mai central . „ .. , ,. . . „ 

d ' ,- , ,.. ; i,.,,„..o JH.1* numerous dorsa lv extending almost to arm tips, ataoffl 

tuanules and radiating scale-like pciiphetal granules >*"< • & i 

R i ■-- 2.65*0.06 {±95% el) N. occllijera intermargmally and actmally. Pediecllariae absent. 

— Marginal plates with «>apr8 granules or granules similar Colour red or orange according to Clark (1946), or pale 

to aclinal surface; tabulae with central granules rarely pj,^ t0 p a | c orange-pink (L. M. Marsh, pcrs. eomm.). 

flat, peripheral granules wedge-shaped sometimes fe ma rk£ The form and granulation of the tabulae 

'f f ^"^^ dih|ribution V (he papulat , ll( , lm c 

tabulae with low Hat central granules, then peripherals y £• 

not radiating and R/r 3.11 i 0.12 (±«M e.l.) . 7 granulation of the marginal plales and the relationship 

7 Tabulae with peripheral and central granules few in of R/r gives this species a particularly characteristic 

number (usually each < 20), irregular in sue and .shape, appearance. The relationship of this species with 

round in cmss-seeiion, peripherals radioing; dorsal arm cxtremc [orrns ^ ft &&!&& from the Great Australian 

plates alwavs indistinct distally with enlarged central „. • , , . '.-. .^.- 

Ettl* " V w/som B.ght arc discussed under that species. 

Tabulae numerous peripheral and central granules On the basts ol a poorly preserved specimen (WAM; 

(uMially >20, often >30), usually regular in sue and 9676, R/r = 35/!4 mni) from Bald l„ near Albany, VV.A. 

shape, peripherals forming cornpaci ring aiouud j n 51.2-64 m, which we recognise as ocelli/era, we extend 

centrals, rarely radialing; central granules convex, (hc pange o| - thc spec j es l0 L | 1C s0l( ,|, c0a *t Q f VV.A., 

usually crowded, larger lhan per.pherals dorsal am, specimens have been collected 

plales with more or less distinct lunils disiaiK and with vhw^vb" * h 

ceDiral granules not prominent (FORM I); eeniral between Geraldton and Cape Naturabsie, W,A., in 

granules irregular m sue, dorsal arm plates indistinct depths of 45-180 in. The specimen from Bald I. was 

distally with central granule or granules prominem (as identified by Shepherd (1967) as N. oceltata. 

in iHjbtvH) (FORM 2); eeniral granules lowei than Distribution: Geraldton ro Bald U near Albany, 

peripher.'ils, becoming flai in extreme cases (as in ft . „. - . 

ocelli/em), dorsal ami plates with disimct limits distally VV ' A " +>~** u "' l«p- »/* 
and wilh even granulation (FORM fc) W oeeltatu 

Nectria ocetlifera (Lauuuck) 
(Figs 1 & 4a) 

Asterias ocelli/era Umarck, 1816: 553; 1840: 239 (part); 

Oudart, 1827; plate I. 
Goniodiscus ocelhjerus: Miiller & Troschcl, 1842: 60. 
Nectria occliifera: Dujardin & Hupe, 1862: 406; Pcrricr, 

1869: 283; 1875; 187; 1876; 3; H. L. Clark, 1914: 1.39; 

1916: 35; 1928; 379 (key); 1946: 86; Shepherd, 1967: 

464, tig. 3 (pari); Marsh, 1976: 217 (table). 
Nectria ocellata: Shepherd, 1967.- 467 (part). 

Material examined: Holotype, MNHN; ECAS 670 
(R/r= 55/20 mm), mers australes?, Peron et Lestieur, 
1803 (Shepherd (1967) has suggested ibat Ihe type 
probably came from C.eographe Bay, W.A.J. 

In addition 65 specimens from WAM, 3 specimens 
from AM and one specimen from MCZ. 

Diagnosis; K up to 75 mm, r up to 30 mm. 
R/'r^ 2.65i 0.06 (±95« / n c.l.). Tabulae with Central 
granules irregular in size and shape. Hat, very low in 
profile (appmx. 0.2 mm above rabula). slightly spaced, 

Necfria oceltata Perriei 

(FigS 2a ii4b-8a) 

Asterias ocelli/era I.amarck, 1816: 553 (pari); 1840: 139 

Nectria oculifent (lapsus eal. for oreltifera): Gray, 1 840: 

Necfria ocellijera. Gray, 1866: 15; Sladen, 1KM>: 3iu, 

pi. 55, figs 1-7; H. L. Clark. 1909; 529. 
Nectria oceltata: K^rricr, 1875: 188; 1876: 4; H L. 

Clark, 1916; 34; 1928: 378 (part), .^79 (key); 1938; 78, 

1946: 85; Cotlon & Godfrey. U>42; 197 (part); A M 

Clark, 1966: 313. 315 (table-part), pi. 2, fig, 3; 

Shepherd, 1967: 468 (part), lie. 4. 1968: 738 (part J; 

Marsh, 1976: 217 (lablc); Zeidler A Shepherd, 1982: 

406 (part); fig. 10.4a. 
Nectria nwlfispinu; Shepherd 1967: 468 (pati). 

Material Examined: Lectotypc (designated herein), 
BMNH; 1958: 7.30.20 (R/r-68/23 mm), tasmania, 
purchased F Germrd jun. One specimen BMNH, 1953, 
4.27.24 (R/r -53-56/16 mm), without data. 

In addition 98 .specimens from AM, 38 specimens 
from MV. 36 specimens from TM, 18 specimens Horn 


REC S. AUST. MUS. J9<9): 117-1*8 

y«wtt wa 

SAM, 15 specimens from MCZ, 7 specimens from 
QVM and 4 specimens from WAM, 

Diagnosis: R up to 125 mm, r up to 45 mm. 
R/r = 2.97±O.05 (±95% elf Tabulae with central 
granules of similar size and shape, close bui usually 
not compact, rounded to angular in cross-section, 
rarely lower than peripherals, Peripheral granules of 
•similar siz.e to each other, usually smaller than centrals, 
wedge-shaped, forming more or less compact ring 
around centrals, tending to radiate in some specimens 
Tabulae varying from flat to convex, extending beyond 
disc to about l A R. Dorsal arm plates usually convex 
sometimes Hat, decreasing regularly in size towards arm 
lip, peripheral granules becoming indistinct, centra] 
granules often enlarged and prominent giving plates 
rough appearance, limits of plates varying from distinct 
to indistinct to naked eye. Marginal pJates distinct, 
granules of similar si?eto each other, convex, close but 
nor compact, similar to those on actinal surface, 
Actinal plates up to 8 rows proximally. Furrow spines 
2-4 (rarely 5). Subambulaeral spines 1-3 on first and 
sceoncl row. Papulae numerous dorsally extending 
almost to arm tips, absent intermarginally and 
actinally. Pedicellariae with 3-4 (rarely 2 or 5) usually 
elongate valves (Hg. 2a), common on all surfaces, 
frequently replacing .subambulaeral spines. Colour 
uniformly brick red to orange, sometimes mottled with 
lighter and darker coloured tabulae. 

Remarks; This is the most variable and frequently 
mis-identified species of Neciria. This has been so since 
Gray (J840) originally mis-identified the species as- 
ocelli/era Lamarck. Perner (1875) examined a number 
of specimens in the BMNH, all apparently from 
northern Tas. and concluded that Gray mis-identified 
ocellifera of Lamarck and proposed the name ocellata. 
We have examined 3 historical specimens each of which 
was possibly examined by Gray and subsequently 
Perrier (A. M. Clark, pers. comm.), of these one 
specimen (BMNH; 1846.8.3.14) we identify as M 
pedicelligera (I mo. 9a) (see p. 000), of the other two, 
one (BMNH; 1958.7.30.20) has tabula and tabular 
granulation conforming closely to Pcrricr's description 
and this specimen we nominate as a lectotype for the 
species ocellata (Fig. 4b). We have rejected the second 
specimen (BMNH; 1953.4.2724) (Fig. 6b) as a possible 
lectotype or paralectotype, even though Perner 
probably saw it, as the tabulae and tabular granulation 
does not conform to Perrier's description of N. 
ocrilata t and we are therefore uncertain thai it qualifies 
as a type specimen. Perrier (1S75) also refers to a poor 
specimen in the MNHN which he was unable to 
determine. This specimen (EC AS 668) which may have 
been seen by Lamarck and identified as ocellifera has 
been labelled as a type of ocellata which it clearly is 
not as the species description of ocellata was based on. 
the material in the BMNH. 

Following our examination. o\' 218 specimens from 
southern and south-eastern Australia w^ have recog- 

nised 3 more or less intergradmg forms within I he 
species ocellata. 

The first form conforms most closely with the 
lectotype in that the tabulae are slightly convex and 
covered with even sized granules (Figs 5a, b), This form 
has its centre o( distribution at the eastern and north 
coast of Tas. extending north to Port Jackson, N.S.W 
and west to Phillip 1. Vic. and is found in very shallow 
water from the shoreline down to 20 m (Fig. 3b). 

Along the east coast o^ Australia we recognise a 
second form with tabulae with central granules which 
become shorter than the peripheral granules, the latter 
radiating giving the tabula a Halter appearance and 
with dorsal arm plates which frequently possess 
enlarged central granules giving the arms a rough 
texture (FlgS 5c, 6a) similar to that of witeoni from 
W.A. This form is distributed between Broughton I., 
off Port Stephens, N.S.W. and Frith L, Bass Strait, but 
has a greater depth range than form 1 occurring in 0-90 
rn (Fig. 3b). 

The third form (Figs fib-d; 7a) has tabulae with 
central granules distinctly shorter than the radiating 
peripherals, becoming flattened and ocellifera -like at 
the extreme western end of their range (Figs 7b; 8a). 
The dorsal arm plates have an even covering of granules 
and the limits of the plates are relatively distinct. We 
do not identify the specimens from the Great 
Australian Bight with ocellifera because of the elongate 
arms and the smaller size and distribution of the 
tabulae despite the fact that the marginal granulation 
is finer than the actinal granulation These extreme 
variants may point to a relationship between ocellata 
and ocellifera which is not obvious when comparing 
ocellijera from W.A. (Fig. 4a) with typical ocellata from 
Tas. (Fig. 5a). The relationship between ocellata and 
ocellifera can only be resolved satisfactorily when more 
specimens from the Great Australian Bight become 

Form 3 extends from south-eastern Tas. west to (he 
Great Australian Bight (125°30'E) occurring in depths 
of 5-230 m (Fig. 3b). 

Although form 1 clearly intergrades with form 2 
along the east coast and with form 3 along the south 
coast, forms 2 and 3 are easily separable and may have 
originated as a result of the Pleistocene land bridge 
between Vic. and Tas, separating the two populations 
which then developed their different characteristics. 

Along the NSW coast, at least as far south as Fden, 
AC ocellata is the only known species oi Nectria so that 
its identity has not been confused. However, along the 
south coast the species most often confused with 
ocellata is peJicelligera (Figs 9b, tOa) which most 
resembles our form 3 and Can mosi easily be 
distinguished from it by the presence of intermarginal 

Of the first 13 records of ocellata listed by Clark 
(1966, table 3) at least two (BMNH; 1862.7.9.51 and 
-52) from W.A. and Dirk Hartog I. (Shark Bay, W.A.) 



respectively arc unlikely lo be oceltata. Further we 
doubt die likelihood of Dirk Hartog I. as a loeaJiry 
from which Nectria would have been collected. 

Shepherd (1967) records a juvenile specimen from 
Caloundra, Queensland; we have examined this 
specimen and confirm its idcniiiy but consider the 
locality data to be erroneous. 

Disfnlnttion; Western end of (he Great Australian 
Bight (I25 D 30'E) through Bass Strait south along the 
eastern co^st of Tas. and north along Hie N.SW coast 
to Broughton I.; intertidal to 230 m (Fig. 3b). 

Nearia macrobrachia H. I . Clark 
(Figs 2b & 8b, c) 

Nectria macrohrachia H. L. Clark, 1923: 236* pi 13, 
figs 5-6; 1928: 379 (key); 1946: 86; A. M. Clark, 1966: 
311, pi. 3, figs 2-3, text fig. 2; Shepherd. 1967- 474, 
fig. 2 (pan); 1968: 738; Marsh, 1976: 217 (table); 
Zeidler & Shepherd, 1982: 406, fig 10.4e. 
Material Examined: Holotype, BMNH, 1929: 6.12 1 
(R/r-60/17 mm), Pelsart Group, Houtman Abrolhos, 
shore. Prof. Dakin. 

In addition 32 specimens from WAM, 23 specimens 
from SAM and 20 specimens from A.M. 

Diagnosis: R up to 75 mm, r up to 21 mm. 
R/r-3.47±0.08 (±95% c,l ). Tabulae with central 
granules irregular in size and shape, flat, low in profile, 
close but nor compact, angular in cross-section. 
Peripheral granules of similar size to each other, 
conspicuously larger than centrals, sometimes radiating 
particularly in deep-water specimens (Fig, Sc), generally 
wedge-shaped, raised well above centrals to form 
concave tabulae. Tabulae low, crowded extending 
beyond disc to about 3d R. Dorsal arm plates flat, 
decreasing regularly in size towards arm tips, peripheral 
granules becoming indistinct from centrals, limits of 
plates indistinct to naked eye. Marginal plates relatively 
indistinct due to close, even covering of granules, 
similar to those on actinal surface. Actinal plates up 
to 3 (rarely 4) rows proximally. Furrow spines 2-3 
Subambulacral spines 2-3 m first and second row. 
Papulae numerous dorsally extending to arm lips also 
present intcrmarginally and actinally. Pedicctlariae with 
3-4 broad valves (Fig. 2b) resembling those of N. 
mnltispina, rarely present, occurring at least on actinal 
and abactinal surface. Colour light yellow r to orange, 
brown or even pink with peripheral granules on tabulae 
lighter and arm tips darker than rest of body (Shepherd 

Remarks: The form and granulation of the tabulae 
(Figs 8b, c) and the distribution of the papulae easily 
separates this species from others in the genus. 

Distribution; Houtman Abrolhos, W.A. to WilsotVs 
Promontory, Vic. and Erith I., Bass Strait, but has not 
been recorded from Tas.; intertial lo ISO m (Fig. 3c). 
Wc have been unable to substantiate the depth record 
of 350 m given by Shepherd (1967) which we consider 
to be a misprint 

Nectria pedieelligera Mortensen 
(Figs 2c r 9 & 10a, b) 

PChaetaster mtwittts, Mcibius, 1859; 3, pi. I, figs 1-2. 
Nectria sp. Fisher, 1911: 163; H. L. Clark, 1914: 140. 
Nectria pedieeltizera Mortensen, 1925: 291, pi. 13, figs 

5-6, text fig. 9; H. L. Clark, 1928: 370 (key); A. M. 

Clark, 1966: 315 (table); Shepherd, 1967; 478. 
Nectria ocel/ata; H. L. Clark, 1928; 37ft 379 (key); A, 

M. Clark, 1966: 315 (table-part), pi. 2, fig. 4; 

Shepherd, 1967: 465 (part); Zeidler & Shepherd 

1982: 406 (pari); fig. 10.4b. 
Nectria multispina: Cotton & Godfrey, 1942: 197; 

A M Clark, 1966: 314 (part); Shepherd. 1967: 468 

(part), fig. 3 (part). 

Material Examined: Holotype, Otago Museum; A 
53-47 (R/r -53/18 mm), Gisborne, east coast North I., 
New Zealand, H. Suter? (see remarks). One specimen, 
BMNH; IS46: 8.3.14 <R/r = 55/18 mm), Flinders I., 
Tasmania, Lord Stanley. 

In addition 146 specimens Irom SAM 3 26 specimens 
from MV, 25 specimens from AM, H specimens from 
WAM, 4 specimens from MCZ and 2 specimens from 
TM and QVM. 

Diagnosis: R up to 120 mm, r up to 36 mm. 
R/r=3T6±0.04 (±95% c.l.). Tabulae with central 
granules of similar size and shapes close or eompacL 
angular in cross-section, larger than peripherals- 
Peripheral granules of similar size to each other, wedge- 
shaped, forming a more or less compact ring around 
centrals, rarely radiating. Tabulae slightly convex, rarely 
flat, extending beyond disc for up to Vi R. Dorsal arm 
plates more or less flat, decreasing regularly in size 
towards arm tip, peripheral granules becoming 
indistinct, central granules similar to peripherals or 
enlarged and prominent giving plates rough 
appearance, limits of plates indistinct to naked eye. 
Marginal plates distinct, granules of similar size to each 
other, convex, close but not compact, coarse, similar 
to those on actinal surface. Actinal plates up to 7 rows 
proximally, coarsely granulated. Furrow spines 3-4 
(sometimes 5, rarely 6). Subambulacral spines 2-3 on 
first row and 3-4 (rarely up to 6) on second row. 
Papulae numerous dorsally extending almost to arm 
tips, also present intermarginally but rarely actinally. 
Pcdieellariae with 3-5 (rarely 6) very slender valves 
(Fig. 2c), valves particularly slender on actinal surface, 
usually common on all surfaces and often replacing 
a subambulacral spine on the adambulacral plates. 
Colour uniformly brick red to orange, often mottled 
with lighter coloured tabulae, especially at the base of 
the arms. 

Kemurks: This is the first time that this species has 
been fully recognised since Mortensen (1925) described 
it, from Gisborne, New Zealand. However, according 
to A N. Baker (pers. comm ), despite extensive* 
collecting efforts in New Zealand particularly off 
Gisborne-, no species of Nectria have ever been 


REC. S. AUST. MUS. 19(9): 117 138 

w> rtwj 

recovered. We therefore believe the New Zealand record 
to be an error and we recognise il as an endemic 
southern Australian species. This species is now known 
to range from Denmark, WA. to Eden, N.S.W. 
including the north coast of Tas. and is a shallow-water 
species occurring in 0-20 m. 

This species has been confused most frequently with 
oceilata and multispina. Thus confusion has probably 
arisen because previous authors have not examined the 
type of pedicel libera thus not appreciating it as a 
southern Australian species believing it to be endemic 
to New Zealand, N. pedicelligera is distinguished from 
oceilata by the intcrmarginal papulae, often the 
presence of a rosette of A prominent tabulae at the base 
of the arms and the more regular form of the 
granulation (Figs 9a, e; 10a). Superficially it resembles 
some specimens of multispina (Fig. 10b) however, 
multispina differs in the form of its granulation, the 
enlarged plates extending along the arm, the 
consistently higher number of furrow spines and the 
much broader valved pediceflariae. 

We have examined the specimen from Westernport, 
Vic. (MCZ, 1932), described by Fisher (1911) and 
examined by Clark (J9I4) and have identified it as 
pedicelligera, The holotype of Chaetasier munitus 
Mobius 1859 is presumed lost as it is not in the Kiel 
museum (Shepherd 1967) and was not found by one 
o\ us (F.W.E.R.) in the Hamburg museum. However, 
the figure given by Mobius (1859) is most like 
pedicelligera, particularly in the arrangement of the 
tabulae and their granulation. The apparent absence 
of pcdicellariae with large valves on the actinal surface 
also excludes it from multispina with which Clark 
(1966) and Shepherd (1967) thought it was eonspecifie 
In order to maintain stability and due To the inadequate 
description given by Mobius and the loss of type 
material we agree with Clark (1966) that the name 
Chaetasier munitus be treated as a nomen oblitum, 
such action however requires a case to be put to the 
ICZN (article 23bl. 

The three specimens (BMNH; 1862.I.8J0 & 11; 
1846.8.3.14) recorded by A. M. Clark from Tas. (1966. 
table 3) as ''intermediate between oceilata and 
muluspina" we identify as pedicel ft gera, 

Distribution. Denmark, WA. to Eden, N.S.W. 
including Bass Strait and the north-west of F&s., 
intertidal to 20 m (Fig. 3d). 

Nectria multispina H. ' , Clark 
(Figs 2d, 10c & 11a) 

Nectria tmdtispina H. U Clark, 3928:375, figs llla-b: 
I93S: 77, 1946: 86; Cotton & Godfrey, 1942: 197 (part); 
A. M Clark, 1966; 314, 315 (table-part), pi. 2, figs 
1-2; Shepherd, 1967; 468 (pari), 1968: 739; Marsh, 
1976; 217 (table), Zeidler & Shepherd, 1982; 406, figs 
10.4c. d. 

Material Examined: Holorype, SAM; K50 
(R/r- 80-85/30-32 mm)? S.A., probably Spencer Gulf 
or Gulf St Vincent, H. L, Clark (1928); 2 paratypes, 
SAM; K52 (R/r -56/19, 51/17 mm) without locality 
data, J. C. Verco Feb 1891; paratope MCZ 2904 
(R/r =60/19 mm) with same data as K52. 

In addition 22 specimens from SAM, 7 specimens 
from AM, 4 specimens from WAM and one specimen 
from MV. 

Diagnosis; R up to 102 mm, r up to 34 mm. 
R/r = 3.0l±0.08 (±95% e.I.). Tabulae with central 
granules irregular in size with central ones being larger 
than those at periphery, very compact giving central 
tabulae smooth convex appearance, angular in cross- 
section, all larger than peripherals. Peripheral granules 
of similar size to each other forming distinct compact 
ring around centrals. Tabulae appear to extend to arm 
tips but from 2 /? R become indistinguishable from 
convex plates, interspersed with smaller, flatter plates. 
Dorsal arm plates with granules maintaining their 
relative sizes but peripherals becoming less distinct than 
on disc, limits of plates distinct ro naked eye. Marginal 
plates distinct, granules often similar in size, sometimes 
the peripherals are noticeably smaller, close, sometimes 
compact, similar in si/c to those on actjnal surface. 
Actinal plates up to 6 rows proximally. Furrow spines 
4-6 (rarely 3 or 7). Subambulaeral spines 3 (rarely 4) 
in first row, 3-8 in second row. Papulae numerous 
dorsally, extending to arm tips, also present 
intcrmarginally; absent aeiinally. Ptdicellaraic with 3-4 
(rarely 2) distinctive broad valves (Fig. 2d), common 
on all sut faces but particularly characteristic of first 
actinal row o\' plates. Colour orange to brick red, 
mottled with darker tabulae, often only some of the 
central granules of the tabulae are darker. 

Remarks: This is a clearly defined species easily 
distinguished by the form of the tabulae, the distinctive 
pcdicellariae and the consistently high number of 
furrow spines (Figs 10c; lla). Although nudttspina has 
not been confused with other species it is clear that 
other species have been confused with multispina due 
to a lack of understanding of the limits of each of the 
species of Nectria, N, oceilata and N. petlicelligera have 
been the species most frequently confused with 
nndtispma and this is discussed under those species. 

Of the 9 lots of specimens referred to multispina by 
Clark (1966, table 3) only the type specimens and 
specimens under the number BMNH; 1962.4.97 are 

The -species which most resembles and appears most 
closely related to multispina h saoria, particularly in 
the arrangement o^ the tabulae and dorsal arm plates. 
However it is clearly distinguished by its smaller disc, 
the consistently fewer furrow spines and the absence 
of the characteristically large pcdicellariae found in 

Distribution: Shoal water Bay, near Cape Peron. 
south off rcmamle, W.A. ro Wilsons Promontory. Vic. 

\U VISION Ul Till C1CNUS \tctKL\ 


bin not from Bass Strait or Tas., intertidal to 20 m 
(fig, 3a». 

j\ectiia nihoni Shepherd & Jfodgkin 
(Figs 2e. lib & 12) 

Necirta iv/7wm Shepherd & Hodgkin* l%5: 119, Rgi 1; 

Shepherd, 1967: 474; Marsh, 1976: 217 (tabic); 

Zeuiter & Shepherd. 198$ 408, fig. I0.4e, 
Nutria oeellattv A. M. Clark, 1966; 315 (table-part), 

pi. L, Tigs 4-6; Shepherd, 1967- 465 (part). 

Ma ferial Exaia ined* ! I o 1 o t y pc, WA M ; 3-65 
(R/l - 77/26 mm), Sorrento Beach. W.A. 1.8 m. B. R. 
Wilson, 27. LI963; paratype, WAM; 18-59 (R/r-63/ 
20 nun), Eagle Bay, Cape Natural iue, W.A., 9.2 m, B. 
R. Wilson & L Marsh, 27.XIf.195S; paratype, WAM; 
19-59 (R/r- 75/25 mm), Dunsborough, W.A., 9.2 m, 
B. R. Wilson & L Marsh, 25. XII 1958; paratype, 
WAM; 2-62 (R/i =88/31 mm), Hatnelin Bay near Cape 
Lccuwin, WA. on jetty piles, B. R. Wilson & R. Slack- 
Sid i t b , 30, X 1 1 . 1 96 U pa ra l y pc W A M , 1 -65 
(R/r- 56/19 mm), Dunsborough, W.A., 16.9 m on 
rock, B. R. Wilson, 15 IV 1963; paraiype, WAM; 2-65 
(R i* = 90/34 mm) Sorrento Reef near Iremantle, W.A., 
on sea grass, B R Wilson, paraiype, WAM; 4-65 
(R/r-90/30 mm) Sorrento Reef, W.A., B. R. Wilson, 
2I.XII. 1963; paratype, WAM; 5-65 (R/r -83/27 mm) 
Sorrento beach, W.A,, 5,5, m on sea grass near 
limestone, & R. Wilson. 23.XI 1963; paratype, WAM; 
3 62 (R/r =120/45 mm), off Beagle Island (29 50'S), 
W.A., Poole Brothers, Mar. 1961; paraiype, SAM; K613 
(R/r-69/22 mm) Hall Bank near Frcmantle, W.A., 
B. R. Wilson, I. II. 1963. 

In addition 40 specimens from WAM, 9 specimens 
from SAM and 7 specimens from AM. 

Diagnosis: R up to 120 mm, r up to 40 mm. 
R/r- 3 08+0.06 (±95% el.). Tabulae with central 
granules irregular in si/.e and shape, a mixture of 
granules larger and smaller than peripherals, markedly 
convex, raised well above labula, well spaced (rarely 
crowded), round in cross-section. Peripheral granules 
radiating, irregular in wc and shape, but forming 
distinct peripheral ring. Tabulae more or less confined 
to disc. Dorsal arm plates, flat decreasing in si/e 
towards arm tip, with very coarse granulation, I or 2 
central grannies on each plate often enlarged and 
prominent, limits of plates indistinct to naked eye. 
Marginal plates distinct, granules coarse, spaced, some 
central ones very large similar to dorsal arm plates. 
Actinal plates up to 7-8 rows proximally. Furrow spines 
3-4 (rarely 2). Subambulacral spines 2-3 on first and 
second row. Papulae numerous doTsally extending 
rarely beyond '■: R, absenl inlermatginally and 
acrinally. Pedicellariae with 4-5 (rarely 3, 6 or 7) slender 
valves (Fig 2c), usually present on all surfaces, some 
specimens without. 

Colour deep orange to magenta, papular area lighter 
(Shepherd and Hodgkin 1965). 

Remarks: Despite sonue confusion between Ihis 
species and ocellata the large and irregular granules 1 of 
ihc tabulae, marginals and arm plates particularly make 
wilsoni a very distinctive species (Fig. lib). The 
granulation o\^ this species is very variable (Fig. 12). 
It is interesting to note thai some N.S.W. specimens-of 
(Hrfiata (Fig, 6a) resemble wilsoni in that Ihc granules 
become irregular. However, the tabular granulation is 
rnueh finer and the granules are more numerous in the 
N.S.W specimens so thai these two widely 
geographically separated species are not confused. We 
agree with Shepherd (1967) that the three specimens 
(WAM; 18.59; 19.59 & 2.62) recorded by Clark (1966, 
table 3) as being "affiliated lo ocellaur are wilsoni. 
The species is now extended in range eastward across 
the Great Australian Bight lo Cape Jcrvis, S.A. 

Distribution Beagle I., W.A, lo Cape Jcrvis, S.A.; 
intertidal to 44 m (Fig. 31). 

Nectria saoria Shepherd 
(Figs 21" 6c 13a) 
Neanu saoria Shepherd, 1967: 475, fig. 2 (part); 1968: 
739; Marsh, 1976: 217 (table); Zeidler & Shepherd. 
1982- 408, fig, 
Nectria mutiispina: A. M. Clark, 1966: 315 (table-part). 

Mater in I Exam ined; H o I o l y pe. S A M ; K 670 
(R/r =56/15 mm), submerged limestone reef between 
Wright I. and The Bluff, Encounter Bay, S.A., 10 m, 
S. A. Shepherd, 8VU963; 4 paratypes, SAM; K628 
(R/r = 56/J5, 53/16, 50/13, 47/14 mm), collected with 
type; 3 paratypes, SAM; K627 (R/r = 6!/I6, 52/15, 
40/13 mm), West J., Encounter Bay, S.A., lOrri, S. A. 
Shepherd, 23.VI1 1.1964; paratype, SAM; K656 
(R/r=65/17 mm) collected with type; paratype, SAM; 
KiSS (R/r= 64/20 mm), limestone reef between Thistle 
and Hopkins I., Spencer Gulf, S.A., 10 m., S. A. 
Shepherd, Jan. 1964; paraiype, WAM 8-64 (R/r=60/17 
mm), Hamelin Bay near Cape Feeuwin, W.A.,. B. R. 
Wilson and R. Slack-Smith, 30.XU.6F 

In addition 24 specimens from SAM, 23 specimens 
from WAM, 7 speeimens from MV and 6 specimens 
from AM. 

Diagnosis: R up to 83 mm, r up to 28 mm. 
R/r = 3.49±0.09 (±95% c.I.I- Tabulae with central 
granules similar in si/c, sometimes 1-2 very large 
granules but these usually mixed with smaller ones, 
compact giving tabulae convex appearance, angular in 
cross-seclion, all larger than peripherals. Peripheral 
granules slightly irregular in si/e forming an irregular 
peripheral ring. Tabulae appear lo extend to arm tips 
but from about z > R become indistinguishable from 
convex plates; interspersed with smaller plates. Dorsal 
arm plates with granulation similar to disc but more 
compact giving the larger distal plates a smoother 
aspect, limits of plates distinct to naked eye. Marginal 
plates distinct, granules often similar in size to each 
other, close, sometimes compact, similar in size to those 


REC, ft AUST. MU5. 19(9); 117-138 tl 19$6 

on actinal surface. Actinal plates up lo 2-3 rows 
proximally. Furrow spines 3-4 (rarely 2), Subambulaeral 
spines 2-3 in first row, Z-4 in second row. Papulae 
numerous dorsally, extending 10 arm tips, less common 
and sometimes absent interinarginally, absent actinally. 
Pedieellariae with 3-5 (rarely 2 or 6) elongate valves 
(Fig. 21), common abaelinally, less common or absent 
marginally and actinally. Colour bright red, papulae 
black (Shepherd 1967). 

Remurks: This distinctive species (Fig. 13a) has been 
adequately described by Shepherd (1967) and its 
relationship to muliispina has been discussed under 
lhal species. Prior to Shepherd's description of this 
species it has been identified with multtipmtL 

OF (lie 9 lots of specimens referred to ntultispitta by 
Clark (l%6» table 3) i hose from Rottnest I., Albany and 
Recherche Archipelago are sctoria as suggested by 
Shepherd (1967). 

Distribution: Off Perth and Rottnest L, W.A. to 
Encounter Bay. S.A. intertidal to 25 m (Fig. 3g), 

Nectria humiiis sp.nov 
(Figs 2g; 13b & 14) 

Material Examined: Holotype, TM; H 1476, SO km 
west of Woolnorth, north-west Tas. t 550 m, A. 
McCifford (**Sea Fisheries ), 2S.Il 1979; paratope, FM; 
H 1844, collccied with holotype; 7 paratypes, AM; 
J 18467, labelled "80/130 off vStorm Bay near Famar 
River 25-30 fms" (approv 45-55 m), nn other data. 

Description: The holotype (Fig. 13b) measures 
R/r = 53-58/17.5 mm; br=22 mm. Arms taper regulai ly 
to narrow lip. Abaetinal tabulae low not more than 
1 mm high, very slightly convex, longitudinally ovate 
up to 3.5 x2.9 mm, spaced, arranged in a earinal row 
with Lwo dorsal lateral rows To about \tt R, a third 
dorsal row of plates also extends to about ' : R, beyond 
! -'2 R the tabulae become indistinguishable from arm 
plates, labulae/arm plates decrease regularly in tffeg 
distaliy. Tabulae with central gianulcs of even size, 
convex, low, angular in cross-section, peripheral 
granules marginally smaller than centrals forming 
compact ring around centrals. Marginal plates discincl, 
22-23 supero- and infero-marginal plates with an even 
covering of small granules (appro*. 7 granules/2 mm), 
liner i nan those on actinal phues. Actinal plates of 6 
rows proximally, the first row extends almost to arm 
lip (to 1 7th infero-marginal plate) the second, third and 
fourth row extend to the 5th, 3rd and 2nd infero- 
marginal plates respectively, the fifth and sixth row do 
not extend beyond the 1st inferomargmal plate. Actinal 
plates covered with coarse, spaced granules (approx. 
5 granules/2 nun). Adambulacral plates with 3-4 
furrow spines, where there are 4 the first one is smallest 
otherwise the other 1 are similar in size to each other. 
Subambulaeral spines, 3 on first row, about half-length 
of furrow spines; 3 on second row, barely indistin- 
guishable in size from adjaceni actinal granulation. 

Oral plates with 8 furrow-spines, 4 subambulaeral 
spines and 7-8 low spines/granules on actinal surface 
of plate. Papulae abundant dorsally in discrete groups 
of cN-lo, confined by (he skeletal reticulum, absent 
intermarginally and actinally. Pedieellariae absent- 
Colour unknown (white in alcohol). 

The paratype (Fig- 14) (TIYF H 1S44) measures 
R/r -46 51/17 19 nun; br = 23 mm. Essentially similar 
to holotype differing only in having stouter arms. 

The seven paratypes {AM, J 18467) range in size 
from R /r — 66/18 mm to R/r-48/25.5 rnm and differ 
from the type only in ihe presence of pedieellariae 
which occur on the dorsal and actinal surfaces, but arc 
most common on the adambulacral plates. 
Pedieellariae with 3-4 valves, triangular in shape, about 
twice as lone as wide (Ftg. 2g). 

from the Latin tlumilts - low, referring to the vet \ low 

Remarks: This species stands apart from other 
species of Nectria by Ihe granulation and form and 
arrangement of the dorsal tabulae. The only other 
species with such low labulae is macrobrachia, but in 
this species the tabulae are close-packed and 
transversely hexagonal or ovate, the granulation is quite 
distinctive and the distribution of papulae immediately 
separates the two species. Amongst the species in which 
the papulae are restricted to the dorsal surface, hamiits 
is most closely abated to ocetlttia. However, the tabulae 
of ocellata never become as low, or the granulation 
covering the tabulae as even, -as in humiiis, 

Jt is unfortunate that the data with the 7 purat\pes 
is ambiguous as the Tamar River is in the mid-north 
of Tas. and the only known Storm Bay is that near ihe 
mouth of the Derwent in south-eastern Tasmania 
(Fig 3b). 


In the most recent revision of rhe Asteroidea, Nectria 
bus been allied with Nectriaster in the subfamily 
Nectriinae of the family Gonasteridae. However, we 
have concluded that skeletal features, including 
particularly the calcified form of ihe tulerbrachial 
septum, show that these genera are better placed in the 
family Oreasteridae. We have further concluded that 
Nectria, by virtue of its abaciinal tabulae, stands apart 
from all other trenera in thai family. We prefer lu 
recognise this isolation by maintaining the status a£ the 
subfamily Nectriinae from which Nectriasrct is 
excluded. All other genera within the family 
Oreasteridae arc referred to the new subfamily 

Wiihin the genus Nectria we have found the presence 
or absence of uitei marginal papulae to be a very useful 
and consistent character for all of the species except 
saoria. This character in combination with the form 
of abaciinal and marginal granulation and the form 



of the pedicellariae have enabled us to determine the 
limits of the species. 

We have determined that Nectria is an endemic 
southern Australian genus and that records from Fiji 
(Sladen 1889), New Zealand (Mortensen 1925) and 
Mauritius (MV specimen) are erroneous. 

Regarding the origin of the genus Nectria in 
Australia, lack of fossil material prevents us from 
determining its evolutionary or temporal origin. From 
our knowledge of the species distributions we would 
infer that the genus arose in south-western Australia 
and distributed along the south coast; this is consistent 
with our present knowledge of major ocean currents 
in southern Australia. However, only one species, N 
ocel/ata, has been successful in reaching the N.S.W. 
coast. Considering our interpretation of the complexity 
of this species we conclude that the full potential for 
speciation in the genus Nectria has not yet been 


We would like to thank Miss A. M. Clark, British 
Museum (Natural History), London; Miss A. Green 

and Mrs E. Turner, the Tasmanian Museum and Art 
Gallery, Hobart; Dr A. Guille, Museum National 
d'Histoire Naturelle, Paris; Dr A. C. Harris, Otago 
Museum, Dunedin, New Zealand; Mrs L. M. Marsh, 
Western Australian Museum, Perth and Dr R. 
Woollacott, Museum of Comparative Zoology, 
Harvard, U.S.A. for the loan of material. Collections 
have also been examined in the Museum of Victoria 
and the Queen Victoria Museum and Art Gallery, 
Launceston by the authors access to which was given 
by Sue Boyd and Bob Green who are respectively 

Miss A. M. Clark and Mrs L. M. Marsh also kindly 
reviewed the manuscript and made helpful suggestions. 

The photographs were taken by Miss H. McClelland 
of The Australian Museum, who is duly acknowledged. 
Ms J. M. Thurmer of the South Australian Museum 
prepared Fig. 3. We are especially grateful to Dr D. 
Kuhlmann, Museum fur Naturkunde der Humboldt — 
Universitat, Berlin, DDR, for supplying us with copies 
of Oudart's figure. 

The Mark Mitchell Research Foundation is acknow- 
ledged for granting financial assistance to one of us 
(W.Z.) for travel to Sydney to complete the study. 


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Archipelago during the years 1842-1846, together with an 

excursion into the interior of the eastern part of Java. Vol. II. 

London, pp. 339-354. 
GRAY, J. E. 1866. Synopsis of the species of starfish in the Btitish 
Museum (with figures of some of the new species). London. 

iv + 17 pp., 16 pK. 
LAMARCK, J. B. P. 1816. Histoire naturelle des animaux sans 

vertebres. Ed 1. Vol. 2. Paris. 568 pp. 

LAMARCK, J. B. P. 1840. Histoire naturelle des animaux sans 

vertebres. Ed. 2, Vol. 3 (reviewed and augmented by G. P. 

Dcshaycs and H. Milne-Edwards) Paris. 770 pp. 
MARSH, L. M. 1976. Western Australian Asteroidea since H. L. 

Clark. Tha/assica Jugostavica 12: 213-225, 2 figs, 1 table. 
MOBIUS, K. 1859. Neue seesterne des Hamburger and Kieler 

Museums. Hamburg. 14 pp. 4 pis. 
MORTENSEN, T. 1925. Eehinoderms from New Zealand and the 

Auckland-Campbell Islands. III. Asteroidea, Holothurioidea, 

and Crinoidea. Vidensk. Medd. naturh. Foren. K'bh, 79: 263-420, 

70 figs, 3 pis. 
MULLER, J. and TROSCHEL, E H. 1842. System der Asteriden. 

Braunschweig, xx + 135 pp., 12 pis. 
OUDART, M. P. 1827. Cours d'Histoire Naturelle contenani les 

principales espece du Regne Animal. Engelmann, Paris. 
PERKIER, E. 1869. Reserches sur les pedieellaires et les ambulacres 

des asteries et des oursins. Ann. sci. nat. 12: 197-304, pis 17-18. 
PERRIER, E. 1875. Revision de la collection de stellerides du 

Museum d'Histoire naturelle de Paris. Arch. Zoo/, exp. gen. 4: 

PERRIER, E. 1876. Revision de la collection de stellerides du 

Museum d'Histoire naturelle de Paris. Arch* Zoo/, exp. gen. 5: 

1-104, 209-304. 
PERRIER, E. 1894. Stellerides. Exped scient. Travailleur-Tulismun, 

1-431, 26 pis. 
SHEPHERD, S. A. 1967. A review of the starfish genus Nectria 

(Asteroidea: Goniasteridae). Rec. S. Aust, Mus. 15: 463-482, 

4 figs, 1 table. 
SHEPHERD, S. A. 1968. The shallow-water eehinoderm fauna of 

South Australia. 1. The asteroids. Rec. S. Aust. Mus, 15: 729-756, 

1 fig., 5 tables. 
SHEPHERD, S. A. and HODGK1N, E. P. 1965. A new species of 

Nectria (Asteroidea: Goniasteridae) from Western Australia. J. 

Row Soc. West. Aust. 48: 119-121, 1 fig. 
SLADEN, W. P. 1889. Asteroidea. Rep. Sci. Res. "Challenger" (Zool) 

30: 1-935, 118 pis. 
SPENCER, W. K. and WRIGHT, C. W. 1966. Asterozoans. In 

Moore, R. C. (ed.). Treatise on Palaeontology, Part II. 

Echinodermata 3. Vol. 1. Geological Soc. America Inc. and 

University of Kansas Press: pp. IJ4-U107, figs 1-89. 
VIGUIER, C. 1878. Anatomic compare du squellete des stellerides. 

Arch. Zool. exp. gen. 6: 33-250, pis. 5-16. 
ZEIDLER, W. and SHEPHERD, S. A. 1982. Sea stars (Class 

Asteroidea). In (Shepherd, S. A. and Thomas, I. M., eds). 

"Marine Invertebrates of Southern Australia!' Part 1. Adelaide. 

S.A. pp. 400-418, figs 10-3-10-9, pi. 30-2. 


REC. S. AUST. MUS. 19(9): 117-138 

June, 1986 

FIG. 1. Oudarfs (1827) figure ol N. ocetlijeru Lamarck. 






Abrolhos * Oeratdton 

Wilsons Prom. 
8 ""VErith Is 

Dirk Hartog 1^ 


Wilsons Prom. 

Rottnest Is. 

Encounter Bay 

Beagle Is 

N. »>orl« 

Woolnorthr-<lTamar River 
Storm Bay 

FIG. 3. Maps showing distribution of Nectria species. 


REC. S. AUST. MUS. 19(9): 117-138 

June, 1986 

FIG. 4. (a) N. ocellifera, R/r = 40/15 mm, "Diamantina" Stn. 68 (30°37'S; 114°44'E), 139-146 m. (WAM; 1375-74); (b) N. oceliata, 

R/r -68/23 mm, Tas. (Lectotype, BMNH; 1958: 7.30.20). 



FIG. 5. N, oceUata (a) Form 1, R/r = 123/40 mm, off Maria I., Tas. (TM; H 1293); (b) Form 1, R/r = 76/22, Cape Woolamai, Phillip I.. 
Vic, 10-15 mm (AM; J12499); (c) Form 2, R/r = 88/29 mm, Broughton I. near Port Stephens, N.S.W., 25 m (AM; J 13003). 


REC. S. AUST. MUS. 19(9): 117-138 

June, 1986 

FIG. 6. S. acellaw (a) Form 2, R/r-97/31 mm. off Norah Head. N.S.W. 50-70 rn (AM; J 3466); <b) Form 3, R/r = 56/ 16 mm, without 
data (BMNH; 1953: 4.27.24); (c) Form 3, R'r-87 29 mm, Wcsiernport, North Arm Channel. Vic. (AM; J 9920); <d) Form 3, 
R r-8027 mm. Great Ta\lors Bav, Brum 1., Tas. (TM; H 450). 




7 V. melhtUi, lorm 3 (a) R/r-78/25 mm, Cape Jcrvis, S.A., 46 in (SAM; K. 585); <b) R/r-- 75/26 mm. Great Australian Bight, 

146-230 m (AM: E 3642*. 


REC. S. AUST. MUS. 19(9): 117-138 

June, 1986 

FIG. 8. (a)N. ocellata, Form 3, R/r-85/28 mm, Great Australian Bight, 146-230 m (AM; E 3645); (b) N. macrobrachia, R/r = 69/18 mm, 
Fisherman L, Green Head, W.A., 21 m (WAM; 493-79); (c) N. macrobrachia, R/r = 57/16, off Port Gregory, W.A., 110 m (WAM; 27-71). 



FIG. 9. N. pedice/ligera (a) R/r = 55/18 mm, Flinders I., Tas. (BMNH; 1846: 8.3.14); (b) R/r = 85/28 mm, Cape Jervis, S.A. 
K 596); (c) R/r = 85/25 mm, Doubtful Islands Bay, W.A., 1-2 m (WAM; 367-75). 

10 m (SAM; 


REC. S. AUST. MUS. 19(9): 117-138 

June, 1986 

FIG. 10. (a) S. pedicel/igera, R/r^ 49/15, West 1., Encounter Bay, S.A., 4-5 m (AM; J 7554); (b) N. pediceiligera, R/r- 72/22 mm, Stanley 
I., Denmark. W.A., 12-23 m (AM; J 12709); (c) N. mu/tispina, R/r = 80/28 mm, Shoreham near Melbourne, Vic. (WAM; 1399-74). 



FIG. II. (a) N. muftispina, same specimen as 10 (c); (b) N. witsoni, R/r = 94/33 mm, Sorrento Reef, Perth, W.A., 4-5 m (AM; J 7834). 


REC. S. AUST. MUS. 19(9): 117-138 

June, 1986 

FIG. 12. TV. witsoni (a) R/r = 68/24 mm, Cape Jervis, S.A., 10 m (SAM; K 1758); (b) R/r-94/32 mm, Sorrento Reef, Perth, W.A. 4-5 m 
(Paratype, WAM, 1395-74); (c) R/r = 120/42 mm, off Beagle I., W.A. (Paratype, WAM; 3-62); (d) R/r = 90/33 mm, Sorrento Reef, Perth, 
W.A. (Paratype, WAM; 2-65); (e) R/r = 64/18, off Dunsborough, Geographe Bay, W.A., 20-30 m (WAM; 746-76). 



TIG, 13^ (a) N, sauriu, R/r = 74/21 mm, off Roitnest I., W.A., 25 m (WAM; 33-71); (b) N, humilis, R/r - 53-58/17-5 mm. 80 km W. ut' 

Woolnorth, Tas., 550 m (Hototype, TM; H 1476). 


REC. S. AUST. MUS. 19(9): 117-138 

June, 1986 

FIG. 14. N. humilis, R/r-46-51/17-19 mm, collected with holotype (Paratype, TM; 1844). 



by Karl J. Wittmann 


Distinctive features of the mouth-parts and ventral setae on the telson necessitate the establishment 
of a new genus, Notomysis, for Leptomysis australiensis Tattersall. 




Institut fur allgemeinc Biologic der Universirat Wien, Schwar?$panierstr, 17, A- 1 090 Wien, Austria 

(Manuscript accepted 30 April 1985) 


WITIMANN. K J l'>Nfi- ReOescriptum of ihc mysid crustacean, 
Nototnvsts uusltvliensts iTalletsall) comb, nov: representative 
pf a new gOWtf. fitec. S- Aust, Mus. 1900): 139-143. 

Distinctive Features of the mouth-parts and ventral 
setae on the lelsori necessitate the establishment of a 
new genus, Notomysis, for Leptomysis australietisis 


While preparing a revision of the genus Leptomysis 
it became obvious that L. aastratiensis Tattcrsall has 
several characters which are not Found in any other 
known species. Tattersalfs (1527) description was based 
on material collected by H. M. Hale in late 1926 in Gulf 
St Vincent, South Australia, No other Leptomysis 
species has been Found in Australian waters since then. 
In his description Tattersall states; u This is a character- 
istic species of the genus . . !'; however, he gives no 
description of the mandibles and maxillae which are 
crucial taxonomic characters at the species and genus 

The doubtful status of the species made it necessary 
to examine the type material which is kept at the South 
Australian Museum (Reg. No- C 1617), The material, 
labelled as syntypes, comprises five adult and three 
subadult females, seven adult males, and several parts 
of animals. A male with a body length of 9 mm was 
chosen as leetotype, dissected completely, and mounted 
in Swan medium on slides. Seven further specimens 
collected by H, M. Hale in 1925 at the same location, 
not labelled as syntypes, are in the Tattersall collection, 
British Museum (Natural History) (Reg. Nos 1964.1.2!; 
2257-2275). The examination revealed that the 
description given by Tattersall (1927) is somewhat 
incomplete; also there are several conflicting points 
concerning features common to all specimens studied: 
the eyes are much smaller; the exopod of the fourth 
male pleopod has four instead of five large modified 
setae; the dactylus of the third to eighth thoracic 
endopods is much smaller than described; and the 
telson bears setae and has a minute apical incision. 
These findings show that a redescription us necessary. 
From the following description of mouth-parts and 
telson it becomes clear that the species cannot remain 
in the genus Leptomysis but should be placed in a new 

Notomysis nov. gen. 

Diagnosis: Mysidae, Leptomysini with eyes normal. 
In addition to the usual sexual dimorphism of the 
amennula the males are characterized by a hairy 
rounded organ located dorsally at the distal segment 
of the sympod. Antennal scale setose all around, with 
small terminal joint. Labrum normal, without 
splniform process. Mandible with processus molans 
reduced, palpus normal. Maxillula with endite small, 
distal joint slender. Maxilla with exopod normal, 
eudopod without spines, proximal endite as broad as 
other 2 endues combined. First thoracic endopod 
without endites, ischium distinct but very short. Third 
to eighth thoracic endopod with carpopropodus 
3-scgmented by transverse articulations. Female with 3 
pairs of oostegites. Pleopods of female represented by 
simple setose plates. Pleopods of male all biramous, 
well segmented except first endopod; fourth exopod 
with a total of 4 large modified setae on distal 3 
segments. Endopod of uropod with spines on inner 
margin. Exopod without distal joint and without 
spines. Telson with numerous setae ventrally, minute 
apical incision, and spines on lateral margins, 

Tvpe-speeies: Leptomysis aastraiiensis Tattersall. 

Etymology: From the Greek term "nbtos % ( = south). 

Relationships: The genus Notomysis is closely allied 
to the genus Promysis t sensu \\ (1964), with which it 
shares features of the antennae, mouth-parts, 
thoracopods, and pleopods. Maxilla, maxillula, 
mandible* and first maxilliped are unlike Leptomysis 
but clearly belong to the type represented by the genera 
Promysis and Prionomysis, The new genus is 
distinguished from these and all other genera of the 
family Mysidae by the unique structure of the telson. 

Notomysis aastraiiensis (Tattersall) comb. nov. 
(Figs 1-26) 

Diagnosis: Middle segment of antennular peduncle 
with strong modified seta directed laterad. Inner 
margin of endopod of uropod with 16-28 spines 
increasing in length distally; 11-22 of these spines 
irregularly arranged on statocyst; 5-7 spines linearly 
arranged distal to statocyst. Telson with about 45-60 
closely set spines on each lateral margin; apex with 
minute narrow incision about %\ length of telson. 
Ventrally telson bears 8-12 plumose setae on 


REC. S, AUST. MUS. 19(10): 139 143 

June, 1986 

longitudinal ridge running half-way between centre and 


Description; General body proportions slender, 
closely similar to Prionomysis aspera li. A further 
coincidence with this species is that body and 
appendages are densely covered with small cuticle 
structures (Pig. 5). These are of about cylindrical shape 
on the carapace, body trunk, eye-stalks, and bases of* 
appendages. Towards tips of appendages they tend to 
become acute and scale-like (Figs 3, 14, 19, 21, 23). 
Carapace with a transverse straight row of ca 13 pores 
in median position a short distance anterior to cervical 
sulcus; a further row of ca 27 pores at the cardial sulcus 
and a larger pore surrounded by 7 smaller ones a short 
distance anterior to the rounded posterior margin of 
carapace. Rostrum large, longer than distal segment of 
the antennular peduncle. Eyes as in Fig. 1. Last 
abdominal somite terminally produced into 2 strong 
spiniform projections on each lateral margin. 

Mouth-parts: Frontal border of labrum bluntly 
rounded. Mandibles with processus molaris reduced to 
two lobes, masticatory lamellae completely lacking. 
Teeth of pars incisivus and lacinia mobilis larger on 
left than on right mandible. Left pars incisivus with 
3 large and 3-4 smaller teeth, with the si/e decreasing 
proximally. Right pars incisivus distally with o^q large 
tooth flanked on each side, rostrally and caudatly, by 
one masticatory plate carrying several small teeth. Left 
lacinia mobilis with 5-6, right one with about 4 teeth 
of varying size, Each pars centralis ( = spine row) with 
4-3 teeth of about equal size. Left mandible only 
densely covered with bristles on caudal face in region 
of lacinia mobilis and pars centralis. Palpus with 
unusually dense setation. Maxillula as is typical of 
Mysidopsis and Promysis, distal joint without array of 
pores found in Leptomysis. Maxilla essentially as 
figured by li (1964) for Ptomysis onentalis Dana. 

Thoracopods: First exopod with 9 segments, second 
to seventh with 10« eighth with 9-10. Proximal segment 
Hanked by large intersegmental joints; acutely pointed 
at laterodistal corner; small plumose seta usually 
present close to this corner. Epipodite 1 linguiform, 
without seta. First and second endopod like those in 
Promysis. Merus of first endopod without array of 
pores that occurs in Leptomysis\ propodus and dactyius 
without spines except a strong nail at apex of dactyius. 

Carpopropodus of third to eigluh endopod with 2 short 
joints; dactyius minute with slendeT nail. First pair of 
oostegites reduced, but larger than usual in Mysidac, 
with same basic setation pattern as in second pair (figs 
17, 18). Penis short, with about 11 smooth, curved setae 
around cjaculatory opening, and a series of plumose 
setae along outer margin. 

Plcopods: In females all 5 pairs represented by simple 
setose rods increasing in length caudally. In sub-basal 
position each bears a more or less indistinct apophysis 
directed laterad. Ali features of male pleopods 
essentially as in Pmmysis otienta/is, sensu li (1964). 
First to fifth endopod with I, 9, 9, 9, and 8-9 joints, 
respectively. Basal joints each with a well-developed 
apophysis which is slender in first endopod, but 
rounded, plate-like in second to fifth. Basal segment 
of fifth endopod with small additional apophysis. First 
to fifth exopod with 8, % 10* 11-12, and 9-10 joints, 
respectively. Fourth exopod with large modified setae 
on last 3 joints. Antepenultimate and penultimate 
segment each with one large" modified seta and an 
additional small smooth seta. Distal segment minute, 
with 2 less powerful modified setae. 

Uropods: Fxopod setose all around, 1.6 times as long 
as telson, or 1.5 times endopod. Fndopod with 
spiniform projection dorsally at statocyst. Distance 
between apical spine at inner margin and tip is 25-30% 
length of endopod. 

Nauplioid stage: Antennuia distally with acute scales 
arranged in comb-like units. Antenna and mandible 
smooth. Abdomen covered with small hairs. 

Length: Measured from rostrum to tip of lelson 
excluding spines. Length is 8-13 mm in adult females 
(n=8) and 7-11 mm in adult males (n-9). Mean egg 
diameter is 0,49 mm (n-4), 


I am greatly indebted to Dr Joan Ellis (London) and 
Mr Peter Aerfeldl (Adelaide) far the trouble they have 
taken to send the material to Vienna. 

kkh:ri:n< i:s 

II, N., I%4. Fauna -lnportit_a, Mysidac. 610 pp. BioRcographical 

Socicry ot Japan, Tokyo. 
TAtTliRSALL, W. M. (*>27 Australian opossum Oh imps 

(Mvskluueu). fiiv. S. Ami MWl. 3: 235-257. 



FlOS 1-9. Noiomysis australiensis (latlersall), l.ectotype, rf 9 ram. I. Anterior body region, dorsal view. 2. Antenna, vemral viesft 3. 
Anlennula, dorsal view. 4. Right mandible, caudal view. 5. Examples ot cuticle structures, schematically. 6. Labrum, ventral view. 7. 
Maxillula, caudal view. 8, Maxilla, caudal view, 9. Masticatory margins of mandibles, caudal view; details show dentation ol pars lnctsiv us, 
lacinia mobilis, and pars centralis of left mandible. 


REC. S. AUST. MUS. 19(10): 139-143 

June, 1986 

FIGS 10-16. Noiomysis itustmlicnsis (Taitetsall), Uviotypc. rf ') mm, 10 lirsi ihoraeic sieiiuie with appendage, caudal view. II. Seamd 
rboiacic appendage, mslral view 12. Distal poiiion ol second ihonicic appcndatic, seiae omiiied. 13-15. litsl, lourlh. and liflh plcopod. 
caudal view. Id. Penis. 



r lOS 17-26. Notomysis aastra/iensis (TattersallK 17-20. ? 9 mm. 17. Second opsiegire, inner face. IK. First oosteghe, inner lace. 19. Fifth 
plcopod, rostra! view. 20. First pleopod, rostral view. 21, 22, Leetotvpe, cf 9 mm. 21. Uropods, ventral view. 22. TeUon, ventral view. 
23. ef 8 mm, 'tarsus' of third thoracic endopod, detail shows daetylus with nail, 24-26, Nauplioid larva, lateral \iews. 24. Distal portion 
of abdomen. 25. Nauplius appendages. 26. Distal portion ot amennula. 



NUMBERS 11-14 

MAY 1986 







North Terrace, Adelaide 
South Australia 5000 






A trombidoid mite (Acarina: Trombellidae) larva, parasitic upon field crickets Teleogryllus 
commodus (Walker) in northern New South Wales, has been reared experimentally to the active 
deutonymph. The larva, protonymph and deutonymph are described as Trombella alpha n.sp., and 
the deutonymph compared with previously known species of the genus, particularly T. warregensis 
Hirst and T. adelaideae Womersley from Australia. 





Honorary fo&ead?ll Associate, South Aimraliail Museum, North terrace, Adelaide, Sotnti \usiralia ?oiio 

(Manuscript accepted 23 January 1985) 


S0( I HUM I. K. \\ |gg& On tmtnhi'tia alpha D.5B. ( Varimi: 
Irnmhtlliclae); t'or relation, doc i i pi ion, Jcvelopmenial 
.itiiHinnalitiL's, 'ivsioiiiaiics and possible ami not \ mhu lurt-v Mr. 
s. Ausi. Mus. Vi(\\): 145-168. 

A (rombidotd mile ^carina: Trombcllidac) larva, 
parasitic upon Held crickets TdtQgrythtS commodus 
(Walker) in northern New South Wales, has been reared 
experimentally to the active deulonymph. The larva, 
protonymph and dcutonymph arc described as 
1'mmbellu alpha n.sp., and the deutonymph compared 
with previously known species of I he genus, 
particularly T. warregensis Hirst and 77 adelaideae 
Womersley from Australia. 

One reared nymph of 7? alpha had gtosa 
developmental abnormalities o\ (he legs, wilh 
shortening or even loss of segments, and non- 
development of setae, here termed brachymely. Similar 
occurrences in tlm and other mites are instanced. 

A correlation and a principal component analysis 
was made of I he larval dimensional variates, using 
various dorsal scutum, leg segmental, and body setae 
measurements. Three principal components were 
defined, each including one or more icliosomal vanates 
and one or more leg variates. 

A key is presented lor the known adults and 
deulonymphs of Trombella of the world. 

leg chaetotaxy of Trombelia alpha, the only 
described larva of Trombelia, is discussed. 

I he family Trombcllidac and its subfamilies 
( ln/criinac and Trombelhnae arc redefined, Generic 
classification of the Trombellidae is revised, A new 
genus is creeled for adults and nymphs: 
Maiputrombclla n. gen., with type species A/. 
americanum (Robaux, l%8) its only member, from 
South America. Keys are provided to the genera o\' both 
larvae and of adults and deulonymphs. Womersieyta 
Radford, 1946 is restored 10 full generic status. 

The adult genus Parachvzeria Hirst is removed from 
the family and placed in the Johnstonianidne, as is also 
the laival genus Ralphaudyna Vcrcammen-Graiidjean 
et at. (1974). 

The possible functions o\' some of the structural 
peculiat ities ol Iromhetla alpha larva are discussed in 
relation to existing knowledge and suggestions that 
have been made for some other prostigmatie mites. The 

htm: 1986 

attenuation of the larval tarsi is believed to be related 
to ils function as a sound receplot, serving as a sonar 
device for the location of the sound-emiuing cricket 
hosts. This suggestion is supported by a significant bias 
(over 3:1) in the numbers of mites obtained from 
vocalizing male crickets as against the silent females 
and because this bias was not found in the sub-adult 
(i.e. non-stridulating) male instars. 


Berlese (1887) proposed the genus Trombelia for T 
gianduiosa Berlese, 1887 from northern Italy; a speciev 
since recorded from Austria (Schuster, I960). Originally 
Berlese gave a definition and description only of the 
type species, i.e. by referring to the sixteen dorsal 
depressions. In 1888 he added the South American T 
nothroides Berl, 1888, which lacks (lie founded dorsal 
opisthosomal depressions. 

Womersley (1954b) reviewed the subfamily 
Trombellinac Trior, 1935 and erected the germs 
Nothrotrombidium for Tiombelia otiorum Berlese, 
1902, from liurope (sec Feidcr 1955, p. 68; 1958, p. 265), 
thereby restricting Berlese*s genus; he placed in it also 
T nothroides Berlese, and T lundbladi Willmann, 1939 
from Madeira (North Atlantic region). Additional 
species include Nothrotrombidium bulbiferum 
(Willmann, 1940) {nam. emetul), from Furope and 
Nothrotrombidium brevttarsum Andre, 1960 (num. 
emend.) (1960a) from Tonkin, Indochina. Trombeilas^ 
str. has currently five described species- T. glandulosa 
Berlese (type), T, warre^ensis Hirst, 1929 (Australia), 
T. adetaideae Womersley, 1939 (Australia), L favosa 
Andre, 1936 (1936a) (Africa) and T lustiamcu Andre, 
1944 (Furope). 

Genera now placed in the family Trombcllidac 
include Trombelta Berlese, Parachyz.enu Hirst, 1926 
(however, see further below), Parathrombetia Andre, 
I95K, Neonolhrothromlftdium Robaux. 1968, based 
purely on the adult forms, Chyz.cria Canestrini, 1897 
and Nothrotrombidium Womersley, 1954, known fuuu 
the adult and the larva. Audyana Womersley, 1954 
(1954a) and Ditrenia Vcreammen-Giandjean, 1955, 
known from the larwt and dcutonymph, d\Ml 
Womersieyta Radlord, 1946, Ral(>haudyna Vcieammen 
Grandjean et t//., 1974, and Noihrotrombicuia 
Dumblcton, 1947, known only from the larva. 


REC. S. AUST. MUS. 19(11): 145-168 

June, 1986 

I reported that larval trombidioid mites, parasitic 
upon field crickets, Teleogryllus commodus (Walker), 
in northern New South Wales, and classifiable to 
Womersleyia Radford, had been reared experimentally 
to the nymphal stage, and that these nymphs were 
classifiable as Trombella (Southcott, 1982). It was thus 
proposed that Womersleyia Radford is a junior 
synonym of Trombella Berlese. Previously it was known 
only from its type species, W. minuta Radford, 1946, 
a larva taken either free-living from mud, or parasitic 
under the wings of grasshoppers (Acridoidea: 
Tetrigidae and Tettigoniidae) on the Island of Gan, 
Addu Attoll, Maldive Islands; original specimens taken 
on 20.xii.1944. (A paratype slide in the South Australian 
Museum collection is labelled (in the writing of one 
of Womersley's technical assistants) "ex Grasshopper/ 
Maldive Is/13, Jan 1945 C.D.R." This slide bears also 
a label in pencil showing that it was used by 
Vercammen-Grandjean in 1970 in his revision (1972) 
of the species. I have added an identifying number 
ACB731 to each label.) 

This paper describes the larva, protonymph and 
deutonymph instars of the New South Wales species 
of Trombella, details of the experimental correlation 
and its taxonomic significance. 

The reared deu to nymphs appeared to be 
morphologically similar to the adult Trombella 
adelaideae Womersley, 1939, known from a single 
specimen collected "from under a stone at Burnside", 
in the Adelaide district, South Australia, in August. 
However, there are differences. One other Australian 
species is known, based also on a single specimen; this 
is T. warregensis Hirst, 1929, found "under a log on 
the bank" of the River Warrego, 4 miles west of 
Barringun, New South Wales, in August 1928 by Hirst. 
The total number of species of Trombella in Australia 
is conjectural, as these mites have been collected and 
surveyed very inadequately. I have seen several as yet 
undescribed species of larvae referable to Trombella s. 
str. parasitic upon grasshoppers, whilst surveying the 
mite ectoparasites of grasshoppers in the Australian 
National Insect Collection, CS1RO Division of 
Entomology, Canberra. In view of the uncertainties of 
attempting to correlate deutonymphal and adult 
trombidioid (and other prostigmatic) mites on 
morphological features, it is considered wisest to erect 
a new taxon, Trombella alpha n. sp., for this species. 

After again studying the characters of Trombella 
alpha and other larvae known in the family, 
Womersleyia is restored to full generic status (see 
further below). 

All measurements are given in micrometres (/tm) 
unless otherwise stated. 

Genus Trombella Berlese 


Note: I have not attempted in the synonymic list for 
Trombella below to define where it covered also 

Nothrotrombidium Womersley, 1954. All authors used 
the genus name of Trombella for all species here 
considered until Womersley's action. It may therefore 
be taken that from 1887 to 28 May 1954 (the date of 
publication of Womersley, 1954b) all usages of 
Trombella included, or implied the inclusion of, 
Nothrotrombidium. Subsequently, the great majority 
of authors used this separation, exceptions being Feider 
(1955, 1959a), Daniel (1959) and Krantz (1978), who 
still included T otiorum Berlese, 1902. These remarks 
apply also to the incorrect spelling as Thrombella by 
some authors. Feider subsequently (1958) became aware 
of, and accepted, the use of Nothrotrombidium. 

Trombella Berlese 1887, fasc. 40. no. 2; 1888, p. 180; 
1893, pp. 91-92, 96, 138, 149; 1894, fasc. 72, no. 6; 1902, 
p. 127; 1912, pp. 2, 4, 8, 9, 11, 14-16, 18, 22-30. Hirst, 
1929, p. 168; Vitzthum, 1929, pp. vii, 63; 1931, 3(11), 
p. 146; 1940, pp. 144, 145; 1941, pp. 506, 826. 
Womersley 1934, pp. 181, 185; 1937, pp. 75, 76; 1939, 
p. 149; 1954b, p. 125. Thor 1935, p. 108. Willmann 
1939, p. 15; 1940, p. 215; 1941, p. 59. Thor and 
Willmann 1947, p. 199. Baker and Wharton 1952, p. 
250. Feider, 1950, p. 4; 1955, pp. 26, 41, 68; 1958, p. 
265; 1959b, p. 541. Vercammen-Grandjean 1955, pp. 
253, 260; 1973, p. 109. Vercammen-Grandjean and 
Kolebinova 1968, p. 250. Schuster 1960, p. 5. Krantz 
1978, pp. 278, 351. Southcott 1982, pp. 286, 290. 

Thrombella Andre 1934, p. 472; 1936a, p. 9; 1936b, 
p. 325; 1938a, p. 215; 1944, p. 230; 1958, pp. 14, 15; 
1962, p. 63. Robaux 1967, pp. 3, 4, 7, 109; 1968, p. 453 
(incorrect spelling). 

Trombella alpha n. sp. 

Description of holotype larva (specimen ACB713A; 
also supplemented from other specimens) (Figs. 1A, 
B, 2, 3, 4A, C, 5A, B). 

Colour in life orange. Length of idiosoma (partially 
engorged specimen, slide-mounted) 355, width 215; 
total length of animal from tip of chelicerae to 
posterior pole of idiosoma 445. 

Dorsal scutum trapezoidal, with a broad projecting 
anterior "nasus"; anterolateral angles ("shoulders") 
rounded, obtuse-angled; posterolateral angles evenly 
rounded. Anterolateral margins indented, lateral and 
posterior margins concave. Scutal sensilla placed 
towards rear of scutum, well separated, and closer to 
level of PLs than ALs with only a faint indication of 
setulcs (branches, see Goff et al. 1982) with phase- 
contrast oil immersion microscopy. 

Scutal scobalae (non-sensillary setae) six in number, 
with two AM setae at about middle of "nasus", the ALs 
and PLs towards their respective angles; these setae 
ciliate, slender, tapering, a little blunted, with a single 
large terminal selule in many instances. 

Standard data of the holotype and a series of other 
larvae are summarized in Table 1. Raw data are held 
on file at the South Australian Museum. 



I KL I. Tromheth alpha sp. nov. Holoiype larva. A Dorsal view. 10 scale on left, (legs on left side omitted beyond trochanters.) B. Dorsal 
idiosomal seui. further enlarged (not 10 scale). In this and in subsequent figures; the A sign indicates the seia is shown in noih the 


RfcC S AUST MUS »(.tty 145^8 

June. W6 


DATA OF- ASLklLSOl 1'iAKVAI Dl / » ' Wr/_7 / I |//7f I 
N. SP 

(All mtVlMitVmL'IU-. Ill iHV10IML'Ul>. /ilHI 








■■" ' • 

Sfi fO 





7ft -yo 






- s; x i 


-j 1) 



1 1 




4 Ml 









100 US 




\ r- 




J. 44 










10 14 

. iS 

3 % 



^ - 

:i mi 





is 2J 













: io 








>> j: 




5 1 -60 


: 94 










W 5^ 

M 00 











55 ■*- 






7, ',s.i 

4 W 


* For imiMintim values of DS. 

At times setae are broken in ihe specimens As Table 
I indicates, it is unusual to find ihe BOIted sensillarv 
setae intact In a mounted specimen, During the 
moulring process, seutaJ scobalae commonly fracture. 
A number Of specimens measured here have been 
allowed to moult. This often results in disruption of 
the skin, or its crumpling and folding and usually 
increases the difficulties of measuring. In some cases, 
however, e.g. in the east dorsal scutum, it may result 
in a flattening and better display of the more anterior 
parts of the scutum, as well as better estimates of the 
rectilinear length. In fully or partially kd specimens 
Of the larvae the shield is often, in its anterior part, 
mo curved and obscured for accurate measurements 
on ihe slide, and in such cases estimates of I- and ASB 
tend to be unreliable. 

Eyes 2 + 2, sessile, each lateral pair on a distinct 
ocular plate near PL angle of scutum. Corneae oval, 
anterior with longest diameter 13, posterior with 
longest diameter 17. 

Dorsal idiosomalae slender, ciliate, blunt-ended, 
arising from (he usual small plates or annuli set m the 
cpicuricle, in rows arranged 6. 6. 6, 6, 4, 1; total 29. 

Venter ol idiosoma with a pair of scobalae in area 
between coxae It and III, slender, tapering, pointed, 
ciliate, 41 long. Behind the level of coxae 111 and about 
38 setae, tapering, ciliate, pointed or slightly blunted, 
in irregular transverse rows across the opisthosoma, 
25-35 long. Anus (uroponisauc/.), of two longitudinal 
valves (obscured in holotype; 36 long in ACB712E). 

CoxaIac2, I, 1, normal, pointed, with long setules. 
Medial eoxala 1 placed over about the mid-point ^)i ihe 
anterior coval border, or a little medial to it, 54 long; 
lateral eoxala I placed well laterally, over the heavily 
chilint/ed anterolateral rim of the coxa, 51 long Coxala 
[[ placed towards outer part of coxa, about 1/3 back 
from the anterior border, 47 long. Coxala III placed 

over anterior border, about 1/5 back from its heavily 
chitinizcd and projecting lateral part; 46 long. 

lev MjLinoiiul ha inula 6, 6, 6. l^gs slender, segn'ieiits 
from femur to tibia more or less cylindrical; tarsi 
tapering, attenuated, Leg I 430 long, II 415, III 455 
teach including co.\a and claw ). Pedoeoxal supiaeoxalac 
not identified, presumably absent, 

larsus 1 116 long, by 18 high ai its thickest part, m:ar 
its origin, Til/Gel - 1.33. Tarsus II 118 by 18; 
Tilt/Gel! l.3l.TaiMi.s 1)1 124 l>> -18; lilll/liclll 
1.45. (Tarsal measurements exclude claw and pedicle.) 
1 or uiher leg niririe data, see fable I. One falciform, 
slender claw to each tarsus. 

Leg scobalae (i.e. branched, barbed setae) slender, 
tapering, poinred, rhe setules (barbs) moderately 
outstanding. Leg scobalar formula: trochanters 1, II, 
111 HI), 1(1), 1(1), femora 6-7(5-7), 7(6-7), 6(6), genua 
4(4), 4(4-5), 4(4), ubiae 7(6-9), 6(6-7), 7(7) (figures given 
for holotype, followed by the range observed in at least 
six specimens, in brackets). 

Femora, genua, tibiae and tarsi with specialised setae 
(sec Figs I A, 2). Large striate solcnoidalae arc present 
on leg segments as figured. In addition to these easily 
identifiable setae, the femora, genua and tibiae carry 
small, slender > pointed, smooth i,e. unbranehed setae, 
not optically active, In previous papers I have identified 
these setae by the name of "spinalac". These setae will 
be referred to here as spinofemoralac, spinogenualae 
eic, according to my previous system of nomenclature 
(Southeott 1961a, 1961b, 1963; see also the further 
comments below). Solenotarsalae 1,1, 0. Solenotlbialae 
2(?1), 2(?I), l(?0) (there is *urne difficulty in 
differentiating the more slender ol these setae from 
spinaiae). (A previous statement (Southeott 1982, n. 
317) that the formula for the solenotlbialae in Chyzerio 
is 0, 0, was wrong, it should be 2, 2, 1 as shown in 
the figures to that article.) Vc.siigipubralae L 0, 0. 
Vesligiogenuaiae I, L 0. 

(jiialhosoma small, compact-, the combined chelae 
bases from above almost hemispherical; gnathosoma 
95 long to front o\' cheliceral blades, by 82 wide. 
Cheliceral blades large, transverse, each With thie- 
(range 0-3) relrorse teeth along posterior edge. Antei it*) 
hypostomala pointed, ciliate, 22 long. Palpal coxala 
present, tapering, pointed, ciliate. 36 long. Palpal 
lormula I, 0, 2, 1, 3, 8 or 9. Dorsal palpfemotala 
tapering, ciliate, 35 long, ventral similar bul more 
slender, 35 long. Palpgcnuala tapering, ciliate, placed 
dorsolateral^, 31 long (from ACB71JC; broken in 
Holotype). Palpal tibialae and tarsalae as figured. 
Palpal tibial claw bifid, with the tines short, curved, 
pointed, a little separated, slightly unequal. Palpal 
supracoxala not identified. Galeala absent 

Analysis of ihe Larval Varieties 

In an analysis of the data in Table 1, I have 
eliminated the Sens figures, since only three are 
recorded. Also, as the data for the VarfOUS estimates 



FIG. 2. Trotttbeita alpha sp. noy. Holoiype larva. Ventral view (legs on left side of drawing omitted beyond trochanters). 


RfcC. S. AUST. MUS. 19(11): 145 168 

June, 1986 

for DS are to some extent redundant, these figures have 
been restricted to the maximum of the DS The data 
are thus reduced to a 19 a 12 table of variates. 

A correlation matrix was first calculated from (hese 
figures, followed by a principal componeni analysis 
(Horelhng 1933) In view of (he small number of mites 
only rhe first three (independent) patterns are 

Pattern I; The variates ASB, L, A-P and Til!] (and 
to a less exteat Til] and Gelll) varied in the same way, 
and accounted for about 40*?o oi the patterning. These 
variates refer mainly to estimates of shield length, 
together with that for Tibia HI* 

Pattern 2: The variates AW, P\V, PSB, AM and c iel 
vary together and account for aboul _W<i of Ihe 
patterning. These refer mainly to shield width 
estimates, plus Gel. 

Pattern 3: The variates DS and Til vary together and 
make up about 20% of the patterning. This pattern 
refers only to estimates of the maximum length of the 
dorsal idiosomal setae and of Tibia I, and has no shield 
variatc component. 

li is "micresting char there is in each component a 
representation of a leg variate. It may be commented 
that these findings appear to differ from results 
commonly obtained from rnorphometric studies upon, 
for example, vertebrates, where the first pattern is 
usually of a size variable, li' such were the case here 
one would expect that all shield measurements would 
lend to be represented in Pattern 1. The results could 
be, lo some extent, an indication of the uniformity o^ 
the parasite samples available lor study. 

Description of Protonymph 
(see pp, 160-161) 

Description of Deutonymph 
\\ ie.s 6, 7 A, B, 8 A, B, see also Figs 4C, 0, 7C, D) 

Description based mainly on reared specimen 

Colour in life orange. Idiosoma slender, cordate, 
flattened dorsally. with prominent division between 
propodosoma and rnetapodosoma. Propodosoma 
more or less conical, with base considerably narrower 
at junction with rnetapodosoma. Anterior edge o( 
rnetapodosoma more or less straight, terminating 
laterally in slightly obtuse but rounded shoulders, 
continuing into the posterolateral and posterior margins. 
(In the unfed and unmounted newly emerged nymph 
the posterolateral borders are somewhat concave; see 
Fig. 4D). Length of idiosoma from tip of "nasus" to 
posterior pole 840, greatest width 425, length ot 
propodosoma 200, of rnetapodosoma 640. 

Propodosoma with a pair of dorsal sensillary areas, 
each sensillum being mounted in a smalt boss carrying 
chitinous projections with pointed tips, the whole 
appearance burr-like. Each boss laterally with a pair 

I I- . V }iornb(l(it alpha sp. nov, Larva in situ, parasitic on a cricket 
(SEW by cuLiiicsy oi Mi S I. Davidson,) 

of short, blunted, ciliate setae, 12, 15 long; each ol these 
setae projecting from individual papillae. Anteriorly, 
propodosoma produced into a short blunted point 
overlying chclieeral fangs; from this point the border 
runs back posterolateral^ and almost transversely to 
an obtuse shoulder, and ihen more posteriorad. 

Anterior and peripheral part of dorsal surface ol 
propodosoma with almost smooth 'Year drop" setae, 
each mounted on an individual papilla, an enlarged 
and heightened annulus of the seta. Long axis of 
scobillum of each seta lies more or less transversely 
upon seta shaft, pointing slightly upwards. The more 
anterior of these on propodosoma point more or less 
medially; those on anterolateral parts of dorsum of 
propodosoma point more or less anteriorly (Tig. 7A). 

The more central and posterior pari of propodosoma 
rugose and devoid of setae, the anterior portion, lying 
between the sensilla, being level, the posterior portion 
formed Into two large projecting bosses, with burr-like 
projections of epicuticle; propodosoma bordered by 
'^tear drop** scobalae, and, more laterally, m<m 
attenuated setae. 

Eyes 2 * 2, sessile, lateral upon propodosoma, lateral 
and somewhat posterior to sensilla. Each pair consists 
of an anterolateral and a posterolateral eye, with a thick 
cornea, each about 16 across. 

A single filiform sensillary seta, 96 long, emerges 
from each scnsillarv boss 

Hysterosoma dorsally with 16 oval or circular 
shallow |iiis in two lateral lines, each of six depressions, 
and a median longitudinal row of four. Depressions 
fairly close to each other, and collectively occupy a 
considerable proportion ol dorsum of hysterosoma. 
Anleromosl pit ol median row elliptical, somewhat 
posterior to two anteromost pits of lateral rows; behind 
it arc two circular pits, then an elliptical pit, set among 
i he six most posterior of the lateral pits of the dorsum. 

Each pit of lateral row somewhat elliptical or ovoid, 
second the largest, posteriorad they become 
progressively smaller. 




500 J 


FIG. 4. Trombeifa alpha sp. nov. A Larva in oblique lateral view, undergoing transformation to nymph, specimen ACB712C on 9.iv.l980. 
e eyespot. B Later stage in the larva to nymph transformation, from below, from specimen ACB712B on 9.iv.l980. The nymph is 
developing within the uneast larval skin, two of the larval legs being shown; III indicates larval leg III. In the nymph c indicates 
the developing chelieerae, p the palpi, and 1, 2, 3, 4 indicate the developing nymphat legs. C Cast larval skin and some of the protonymphal 
^kin (latter drawn in heavier line) (specimen ACB712D). The palpi and basis of the gnathosoma are laid back and these parts of 
the mouthparts are seen in ventral view, a the "Y"- or "restructure of the deutonymphal skin, b another part of the deutonymphal 
skin, c- the "matcola" or studded boss of the deutonvmphal skin. D Deutonymph ( ACB712C) seen in transparency. (All to nearest scale). 


REC. S. AUST. MUS. 19(11); 145-168 

June. 1986 





, c 



FIG. 5. Trombella alpha sp. nov., protonymph. A An almost iniael protonymphal skin, from specimen ACB712A. a anal valves from 
the larva, as area sciosa, m mateola or studded boss, n one of Ihe nodular bodies inside the skin (shown cross-hatched), Is attached 
piece of idiosomal skin oT larva, ds dorsal scutum of larva in a fragment of larval skin (to scale on left). B Anal valves and area 
setosa of another specimen (ACB714B) (to scale on right). 



FIG. 6. Tromhettti alpha sp. nov. Active nymph (deuionymph), specimen ACB712B, dorsal view, entire. 

Most of dorsum of hysterosoma densely covered 
with robust setae, with tear-drop shaped scobillum 
perched transversely on chitinized '"papilla" (seta 
annulus). Surface of scobillum with faint more or less 
oblique pattern, visible at high magnification, but 
scobillar surface smooth in lateral view. These setae 
occur over most of dorsum, including pits, but are 
sparser in inner parts of pits. Setae smaller in anterior 
part of dorsum of metapodosoma, and surfaces tend 
to project, thus resembling small burrs; laterally seiae 
tend to elongate and be ciliatc. 

The second, fourth and fifth of lateral row of pits 
contain, more or less centrally, ring of chitin (appears 
as "C" in pit 5) set lower than bases of setae; it possibly 
functions as a respiratory aperture, but could function 
as muscle insertion; presumably derived from seta base 

Ventral surface; not clearly seen in ACB712B, but 
appears to be normal for Trombella. Genital aperture 

and anus appear to be normal for deutonymphs 
(somewhat obscured in mounts). 

Legs (Figs 6, 8A, B) short, fairly robust for a 
trombidioid mite, segments beyond trochanters more 
or less cylindrical. Leg I 620 long, II 415, III 445, IV 
520 (including trochanters to tips of tarsi, without 
claws). Each tarsus bearing two claws. All segments of 
legs with irregular surface due to leg setae (scobalae) 
originating from small papillae. These leg setae 
generally robust, and tend to resemble body setae but 
are much more slender, tapering and blunt-pointed. In 
more proximal and more distal parts of legs these 
scobalae tapering, more slender and Flexible, more like 
usual setation of trombidioid mites. Leg segments 
carrying small spiniform sensory hairs (spinalae) 
interspersed among scobalae, from telo femora to tibiae. 

In leg I (Figs 6, 8A) tarsus from above more or less 
parallel-sided, 198 long by 67 wide. In leg IV (Fig. 6) 


REC. S. AUST MUS, 19(11): 145-168 

June, 1986 



■■ v k« 




9 p 




200 J 





r < 

»ji* « 


, L 


i G 


FIG. 8. Tromhetla alpha sp. nov. Deutonymph. A, B Normal specimen ACB712B, A left leg I, B left leg II. C-F Abnormal legs of specimen 
AC7B7I2A (see te\l), C left leg I, showing extensive deformity, with almost complete loss of all segments beyond the telofemur, but 
all segments somewhat abnormal- D Right lee I, showing a generalized deformity with progressive shortening of segments; note also 
loss Of seiae. E Left leg II, similarly affected. F Right leg II, with considerable deformity, including shortening of all segments and 
gross reduction of setation, and reduction of the paired claws to a single short peg. (All to scale shown.) 

FIG. 7. Jrombelfu alpha sp. nov. Deutonymph. A Dorsal view of anterior part of idiosoma and tips of chelieerae, specimen ACB7I2B. 
B Further enlargement of the setae of the anterior of the median column of depressions. The surface patterning of three of these 
setae is shown at the lower right. C Chelicera ol specimen ACB712C O Medial aspect of R palp of ACB712C. (All to nearest scale.) 


RFC S AUST. MUS. 19(H)- 145-168 

JUlW t /WW 

I Alii 12. MLASUKLMINISUI I 1 IH( - 1 MS him) Ol l.l-G SI.GMI NIS (II A SI KIISOI NYMPHS Ol / KOMM'.U 1 .4/.P//4 






Vromheiia aifihu 

Adult ol 
T Qtieloideav 

Adult ul 

Specimen mirnher 







ALU? 30 



r a i i o 








i alio 





ISrsiia 1 










1 00 





Tibia 1 















Genu I 





— i 










Tarvu- IV 




I 00 






1 00 


! .IK) 


1 1)0 

Tibia IV 


, l >0 













Genu IV 















* Teraiological specimen. The measurements are taken from ilic lets Jelormecl right side (see icmj. LC£S II and III also show the same 

deformity Ibrachymelv). 
I This specimen has a brachymetic righl leg ML However legs 1 and IV appear normal (see teu). 
{ Specimen flexed, not measurable in rnounL 

tarsus tapering, 126 long by 36 wide. For other leg 
dimensions see Table 2. 

Palpi fairly stout, provided with robust scobalacover 
femur and genu, more distally setae more slender, and 
some are barbed (see Fig. 7D, from ACB7I2C). Palpal 
tibia with strong curved claw, and accessory claw-like 
seta on medial side. Palpal tarsus with sparse sctation 
of slendei pointed hairs, some barbed. 

Cheliceral fang robust, curved, pointed, 71 long, 
arising from a basis ehelieerac about twice as long (Fig. 
7C, from ACB7I2C). 

Origin of the Specimens q/'Trombclla alpha 

This study oi' ttomhelta alpha is based upon larvae 
which have been obtained parasitic upon field crickets, 
Teleogryllus cowmocius (Walker), in the New bingland 
District of New South Wales, by Mr S. J. Davidson, 
who commented (pers, comm., 5. si. 1979) on ". . . a red- 
orange coloured mite which appears to be parasitic on 
grytlids. Up to 50 per cricket have been recorded, with 
mouthparts attached to the hosts 1 pleural regions, 
especially on the metathoraV (see Fig. 3). In a further 
note (pers. comm., 1982) Mr Davidson has summarized 
his observations on the parasitisaiion of the crickets 
by this mite species as follows: The mite ''attaches 
principally to the sofl pleural regions o\' I he 
hm( . . , Numbers of this mite were monitored by 
examination of most of the T. eommodus trapped in 
pitfalls from 1979 to 1980, A higher percentage of 
crickets was parasitised in 1979 than in 1980. with 
respective means of 61% and 20% , . • Similarly, there 
was a higher mean parasite burden per cricket in 1979 
than in 1980. The mite was most prevalent on crickets 
in about February, and both percentage parasitism and 
parasite burdens declined in late autumn " [Table 3| 

Material examined 

ACB7I1: 4 specimens, collected from paddock 
number K2, Kirby Rural Research Station, near 
Armidale, N.S.W., I2.ui.l979. Preseived in alcohol, 

\CB7I2: 5 specimens, live, crickets collected 
F2.iii.l980; mites removed 19.iii.1980. (Translnimed in 
nymphs m Adelaide.) 





Number of 

Percentage of 

Mean number 




-.el* with 

t»f miles 





per cricket 









if. i..7y 







2. SO 







0. IS 

Menu (1W) 




1 1 i.MO 












4.iv. 80 




Mean (1980) 




Mole: These data iclci ID all uiMarv of the cricket in which llie *C< 
tail I'l distinguished, i.e. ihHaM ihree instars (see also J ahltf 7, and 

Irxi iheri-liO- 

ACB7I3: Several larvae collected 19.iii.l980; 
preserved in alcohol. 

ACB7M: ? specimens, live, collected I9jii,l980. 

ACB7I2, ACB7I Vand ACB714 were all Collected 10 
km N o\' Armidale, N.S.W. 

Type and paratypes to be deposited in the Sutiih 
Australian Museum. 

Hearing Experiments: Larvae (o Nymphs uj I alpha 

Deutonymphs were reared from two batches al 
larvae forwarded to my laboratory, on strips of wet 
blotting paper in scaled lubes, l:\periiTicuial details are 
as follows: 

(I) Experiment ACB7I2. The cricket was collected flfl 
I2.iii.1980 and ihe mites removed on I9.iii.i980. The 
five larvae sent were received by me o\\ 21. "hi. 1980. 
On receipt the mites were immobile, i.e. they were 
possibly in a pre-pupal condition One mile was 
rather swollen, on its back in a film ol water on I he 
site of the tube. The legs looked decolorized One 
larva was waving its legs about, stuck to glass. The 
three other larvae immobile. flOl swollen Later in 
the day two mites became immobile and were 
considered swollen, while two were waving theit leys 
about AU live were stuck in a film of water and Ihe 
tube was wet. 
23.iii.80. All larvae immobile, plump, 



24.m. 80. AH immobile, not shrivelled, nol mouldy. One 
or more were considered to be in a pre-pupal stage. 

25-2V.iii.80. All miles observed daily, and recorded as 

tO.h1.&0< In Iwo specimens "lioshng" under the skin 
oJ the mile was delected. This was Interpreted as 
'cprcsemint: the developmcnr of nyrnphal setae 
within the exuviae. 

3I.iii.80. 3.00-3.15 p.m. One larva appeared to be sitting 
upon a pupa larger than itself. Skin of larva 
appeared transversely wrinkled. The other larvae 
we a* smooth, .swollen, nor mouldy, and wcic 
considered to be in a pupal state. 
J,3fl p.m. The legs of the nymph were clearly visible. 
Anterior part of the larval skin bad gradually 
become whiter and more opaque, as though Tilling 
with air. A clear gap was visible, most noticeable in 
the right rem leg. 

I he legs of the nymph gradually extended away 
I mm the body o\' the animal. 
Other larvae also had air under the skin. 

I.iv.80. Nymphs had not as yei emerged. Two larvae 
in the lube had prominent protubet'Hnces. Nymphs had still not emerged, despite nymphal 
palpi and legs being clearly visible within the larval 

9.iv.80. Camera Incida sketches were made of the 
transforming larvae (see Figs 4A, B). 

I5.iv.80. Two nymphs had emerged, these being labelled 
ACB7I2C and ACB712LI These were not the 
specimen* mainly described above. The east larval 
skins ol these two mites, also a dead larva 
(ACB712F), were mounted. 

I6.iv.80. One more nymph (ACB712A) emerged in the 
morning, and one (ACB712B) at night. The cast 
skins of these two specimens were mounted. 

IK iv.80. One nymph (ACB7I2D) was immobile, 
possibly dead; mounted. I attempted to feed the 
nymphs with pieces o\' apple* culicinc mosquito eggs, 
aj d '--quashed adults pf Acdes rtotoscripius, also a 
sample of bird dung, and the juice of a squashed 
grasshopper [I J haulacridium yittatum), etc. One 
nymph appeared to feed upon the piece of apple, and 
possibly feeding could have occurred with the other 
m.iiei i;j|s. One nymph (ACB712D) was seen moving 
slowly as late as 17.V.80; the others had died earlier. 
Fvernually all nymphs were mounted in gum chloral 

mountaiU lor study; none had transformed La ' 
later instar, 

(2) Experiment ACB7I4. Three plump larvae were 
received in wet preparation on 2l.iii.S0, having been 
collected on I9.iii.80. All remained immobile, and 
unaffected by mould filaments spreading slowly 
across the inner surface of the glass tube, which 
began to produce spores on 27.iii.80. 

On 28.iii.80 one mite was observed to hove 
decolorized legs and mouthparts. 

On 3 1. iii.80 nymphal legs developing in one larva, 
in its unchanged larval skin, were delected. 

On 8. iv.80 one specimen (ACB7I4A) became 
mouldy. Another specimen (ACB714C) eventually 
shrivelled to a more or less spherical object without 
recognizable features. One larva (ACB714B) was seen 
to have protruding limb masses by 8. iv.80, ajid a 
nvmph emerged on 20.iv.8O. Food was ottered to U 
in the form of a mass of culicinc eggs, pieces of grass, 
a squashed diptcran, a squashed small weevil, a 
squashed Phaulacridittm vittatum etc., and the mite 
possibly fed. Jt was not seen moving after 13.V.1980. 
The mite and its cast skin, and the other larvae used 
in the experiments, were eventually mounted on 
slides for study. 
From these observations a confident correlation of 

the larva with the nymph of this species can be made- 
Table 4 summarizes the details of the successful 

larva-nymph transformations. 

Remarks on the Larva to Nymph Transformation 

The active nymphal stage of the Parasitengona is 
currently recognized as the deutonymph. Between the 
larva and the deutonymph is the protonymph, which, 
as wiih most prosligmatic mites in the Parasitengona, 
is calyptostatic and takes place within the larval skin. 
In the Erythraeoidea the skin tends to retain irs shape 
when shed by the protonymph, and can commonly be 
retrieved in one piece after such ecdysis (Fraucnfeld 1868; 
Womersley and Southcolt 1941; Soul hcott 1946a, 1946e, 
I96la) t In the case of the smaller Trombidioidca ihc 
protonyrnphal skin is thinner and more fragile, so that 
although there have been many studies on larva to 
nymphal transformations in ihe Trombieulidae and 
other families, rarely is there an attempt to describe 
the protonyrnphal stage between the larva and the 
deulonymph. However, modern exceptions to this 
statement can be found in the works of Jones (I95l v 



1 \jH-nmcnl and 



Date became 

Aclive nymph 

Diirali-iti ui ihc 

'.pri'llivrn iu 






immobile phase* 



19-23. iii.80 


24 28 days 

a< k-i:b 


21 .iii R0 




\( in:<_ 


21 iii.SO 

19 -23.iii.80 


JT 21 

\t wrzv 

12 hi. 80 

21 iii.80 

\9~ZJ iii.Kfi 

l c .v 80 



19, iii.80 


19 21 iii.SQ 



• Hv 9Dbl taction, ihe figures given express a mmge ol pov,>ble Jates Irom thr data iwi limle. Since ihtfpe >•< no precise indication ol the 
lime of commencement ot Ihe wnw fi is nol poviihli (o gUtt aflV esiinuU' <«t Uu- duration i>l that period. 


RliC. S. AUST MLS. 19(11): 145-168 

June, ivs6 

1954), Neal and Barnett (l%l), Johnston and Wacker 
(1967) and Robaux (1974), 

An intermediate membrane seen within the investing 
integuments of the prelarva, or later stages, was named 
the "Zwlschenhaut" by Ciaparedc (1868) in his study 
of The development of the water mite Afax bothy 
Ciaparedc, the "apodenna" by Henking (1.SS2) In his 
study of Trotnbutium fuliginosum Hermann, or the 
"intermediate skin" by Jones (1951, 1954), in his study 
on N€otrotttbtrtjfa autuwnttlis (Shaw). Si nee 
deutonyrnphal development takes place entirely within 
the expanding larval skin, there appears no need \'or 
the protonymph of the Trombidioidca to develop the 
distinct pupal setatjon as in the Erythracoidea. and 
which presumably has a delen si a- and possibly 
substrate-anchoring function. 

In the ease of rhe developing Trotnbella alpha 
deutonymph, some of the successive stages of the 
gradual transformation from immobile larva through 
the protonymph to the active deutonymph are shown 
in Fig 4A, Fi Initially the legs and palpi lose the r 
orange coloration, as the living substance of the animal 
is withdrawn to a more compact mass, The ventral part 
Of the larva enlarges, so that after several days Hie 
appearances arc of a larva riding upon a larger rounded 
structure, a flattened prolate spheroid. The uymphal 
leg prominences gradually appear, and project 
increasingly from the main mass. The larval skin 
appears lo separate from the developing nymph, so thai 
the doTsal surface at least appears as whitish and more 
opaque. The emerging nymph may or may not la&ve 
behind a more or less intact larval skin. Generally the 
•intermediate skin" i.e. the protonymphal skin, remains 
inside the cast latval skin, so thai [| is not easy to 
recognize Occasionally parts of the protonymphal skin 
separate from the larval skin, and rarely, the 
protonymphal skin may appear as a more or less intact 
structure. Even within (he cast latval skin, however, 
distinct structures may be seen in the exuviae discarded 
by the nymph. Thus in Pig. 4C there is visible a Y- or 
T-shaped thickening of the integument (V) t shown 
underneath the larval scut urn More posteriorly is a 
distinct structure, near the midline, which appears as 
a studded boss or k *inateola'\ with folds of thin skin 
running up to it </V" in Fig. 4C1 Oilier less clearly 
recognizable structures can also be seen, with vague 
outlines suggesting that they may have been investment 
of the developing limbs, and sometimes with numerous 
small elevations (e.g., as shown in Fig. 4C, '7/ '). 

One distinct structure is made up o( two elongate- 
oval or kidney-shaped valves, surrounded by a large 
number of small conical elevations in an "area sclosa" 
{'as 1 in F ; ig. 5A; see also Fig- 5B). These valves art: ihe 
same Miiuitncas the larval anus or M uroporus", having 
the same appearance of two apposed valves, but 
surrounded by a highly modified area of skin 
Comparison of I he cast larval and protonymphal skins 
shows thai between them they possess only one pair 

of valves or valve-like struct ures, which are of the same 
dimensions as those of the larvae, ix. these are of larval 
origin, the area sctosa, however, is purely of 
proionymphal origin The more or less concentrically 
arranged elevations are short cones with blunted tips. 
' t-v are setiform in shape, and either represent 
precursors CO setae, or possibly serve as investing 
stiuetures for the developing setae of this area of the 
deutonymph. These protonymphal setae are similar in 
appearance to the projections of the 'Mudded boss" 
("niateola'\ or small mace) mentioned above- 

Although it is not at present possible to allocate 
anatomical positions or functions to all of the 
structures of the protonymphal skin, at least some of 
rhe main siructures can be postulated. 

In Fig. 5 A is shown a more or less intact 
protonymphal integument, after slide-mounting 
through lactophenol and gum chloral media (specimen 
AC13712A). This is described briefly in the following 

Description of Casl f'fofutnmpha/ Skirt 
(Fig. >A, B) 

This skin (ACH712B) is more or less in one piece, 
transparent and colourless. Overall length 875, to op 
of projecting anterior pointed structure; width 44s. 
General outline oval, with broad rounded posterior end 
and crumpled but narrower anterior end. Whole of skin 
with crumpled appearance; Only some of the many 
minor folds are shown in Fig. 5A. The appearance of 
the skin matches roughly the shape of developing 
nymph (see Fig. 4B), with widest part and lateral bulges 
eoi responding to legs JV of nymph. The position of 
anal valves and surrounds corresponds to position of 
these structures in larva and deutonymph. The anterior 
part of east skin is more difficult lo assign lo 
corresponding structures, but presumably the various 
parts arc investing integuments roehelicerae, palpi, and 
the more anterior legs. 

The mateola appears in rhe skin, projecting 
anteriorly, about midway between the anal area and 
the front of the skin with folds o( integument leading 
to the base. The u Y-structurc" was not certain I ■. 
identified, bur possibly was contained in the folds 
drawn around the mateola ('>;/" iu Fig. 5A). 

Rounded markings present ovci various parts of cast 
skin, recognisable as rounded elevations on 
protonymphal skin. There is no evidence ol seta 
formation in them (such skin elevations are a well- 
marked feature of the lew developing protonymphs ol 
Irornbidioid mites that have been described; e.g. 
Johnston and Wacker, I%7, Fig. 5, p. 307, for 
iMtmmhiculu splrtvli'iis). 

Several thickened, almost nodular deposits appear 
within the skin, shown cross-hatched in Fig, 5A, one 
being labelled 'V.' Whether these are meie accidental 
inclusions, or are representative of some consistent 


I 59 

process is not known, because only one reasonably 

intact proionymphal skin was avaiiabk. 

Anal valves (Tram larva) and area serosa as 
described above, 

Associated with ihe protonymphal skin in this 
preparation arc only two oilier pieces of larval skin. 
One ("cfe" in Fig. 5 A) consists of the larval dorsal 
senium and a small amount of its surrounding skin, 
detached from the protouymphal skin The seiae of 
scutum are severely damaged from the processes o! 
Iranslorruatiou, and possibly by the mounting 
procedure and no seta is intact. The other small tarvaJ 
skin piece is at the posterolateral aspect of the 
protonymphal skin (V.v' in big. 5A). 

Vunuoons in the l£% Structure oj the Deutonymph, 
and the Developmental Abnormality 'Wrachyntcly" 

Since dimensions oi leg segments are commonly used 
as species -din era i haling characteristics in these and 
other mites, it is instructive to examine the intraspecific 
variation of a batch of mites which have been collected 
from the same area, and handled in the laboratory as 
uniformly as possible. 

bible 2 lists tarsus, tibia and genu lengths of legs 
I and IV ni the reared Trumbcl la alpha nymphs (also 
those ol holotypes of 7. adelaideae and 71 wam^ensis)). 

There is some similarity between ratios of (he 
segmental lengths in the three nymphs ACB7I2B, 
ACB7I2C and ACH7I4R, However, ACB712A shows 
gross shortening ol the three measured segments of Leg 
I, but no abnormality m leg IV. This occurs in greater 
degree distally; and is accompanied by other structural 
abnormalities. It ocairs in legs other than those shown 
in Table 2, and may vary between the two sides of the 
animal. This developmental abnormality will be Termed 

(iraehvtnely in liomhella alpha and in some other 
Frv&tigmutic Miles 

Fig. 8C-F shows the deformed legs I and II of 
AC B712A, Compared with (he normally developed legs 
I and II of ACB7I2B (Fig. SA, B). 

IAUI I. 5. COMPARISONS Ol AHSOI I 1L I £ NO 1 1 IS (urn) Ol 

Mo SEGMENTS oi LEGS I and IV oi klakhd 



M -H7I2A* 



y ■< t • w 











I S3 






















12*. 7 








1 .0 1 




7 C 




* \1c;i-niiM;' the Il*.%\ deformed K teg, tor puipi'\es. of Lomp.tnsnn 
In lh£ Il"i \c& I he $£gfl)ent$ he\onci ihe femur arc unrecognizable 
From tbc ifruirt a&tirttning nad fusion 

' ui J, in Mit can "i Gel 

lable 5 shows normal variation for leg IV in 
specimen ACH7I2A; deformities in this specimen ait 
restricted to legs I-IU. In the right leg 1 the tarsus, tibia 
and genu are considerably shortened, and the setae 
much reduced (Fig. 8D). The trochanters, basil'eniora 
and telolemora appear more or less normal, with a 
minor degree of shortening and some loss of setae in 
these segments also. The degree of shortening is greater 
distally. Ln legs II (Figs. SE, F) and 111 there is a 
reduenon of setae throughout the legs, including the 
two parts of each femur. In right leg JJ (Fig. 8F) and 
right leg 111 there is only one tarsal claw to each tarsus. 
The most extensive delormity occurs in left leg I, when: 
there is some | hickening of the femur, and beyond the 
femur the remainder of the leg is reduced to a single 
fused segment (Fig, SO. In this segment there is no 
recognizable tarsal claw or claws, wirh only a 
rudimentary indication of setae, without any seta basis 
or other pan being distinctly recognizable. (In specimen 
ACB714B there appears to be a minor degree of 
shortening of right leg III, apparently without othci 
abnormalities in the body or other legs. The 
measurements ol the lengths are: Tarsus III 66, tibia 
III 76, genu III 57. In tarsus 111 and tibia III there is 
a reduction of setation, and possibly also in genu IFF 
For left leg 1 1 1 of the same specimen: tarsus III 93 long, 
tibia III 73, genu III 58. 

There was nothing that occurred in the experimental 
handling of specimen ACB712A to suggest a reason 
for the remarkable degree of deformity observed. All 
of the larvae in experiment ACB7I2 underwent 
transformations at about the same time intervals. The 
abnormalities cannot be attributed to a failure of the 
production of moulting fluid, as such would simply 
fail to allow release, but would not be expected to affect 
limb development, Abnormalities other than in the 
limb segments were not detected. The cast skin of the 
larva and protonyrnph appear normal. The lengths of 
the leg segments of the larva, also the shield data (see 
Table I), are normal. The larval tibiae and tarsi, tor 
example, are normal in structure. The cast 
protonymphal skin (see Fig. 5A) appears normal. Jt 
is the most intact of the protonymphal skins available 
for study, at least of those that have separated from 
the larval skin. Jt would be inappropriate to attempt 
to connect some of the parts of the ACB7I2A 
protonymphal skin with structural abnormalities of the 
nymphal legs at the present time> because of the lack 
of knowledge of the correspondence of nymphal and 
proionymphal structures. 

It is not uncommon for minor abnormalities in leg 
daw structure to be seen in larval erythraeid mites 
captured in the field. This has been referred to as 
"twisted claws" or TC abnormality (Southeott, 1961a. 
p. 481; 1966, p. 732; 1972, p. 29). More severe 
deformities arc also seen m the legs of larvae caught 
in the field. Thus in one batch of larvae caughi at 
luinari Valley, South Australia in 1952 many larvae of 


RIT. S. AUST MUS. 19(11); I4&-1M 

./we. /SW<5 

Erythrites urrhrae (Warn.) and E reginae (Hirst) 
showed a mild tarsal deformity, and one specimen of 
£. retinae showed an extreme deformity of all 
pcdolarsi, which tore grossly contracted and distorted 
(Southeott, 1955). 

Ihe cause of deformities is unknown. An 
attempt to reproduce them in larval crythraeid mites 
by chilling hatches of eggs resulted in scuta! deform i tic:-, 
but not leg deformities (Souihcoiu 1955). 

Although it is not unusual for minor abnormalities 
Of scutal structure to occur among prosngmatic mites, 
involving supernumerary setae 01 loss of setae {see 
Soulhcotl, 1966, pp. 756-75S), it appears that Ihe limbs 
of developing mites are more ai risk of structural 
abnormalities (in the form of shortening of segments 
and loss of seurional pattern) than the structures 
associated with the idiosoma. 7 lie claws may suffer a 
minor deformity, may he grossly distorted, or virtually 

Various other developmental abnormalities have 
been recorded among the Acarina and in othci 
arthropods. Among prosiigmatie mites there may be 
compleic absence of a leg, e.g. in the water mite 
Spenhon glandutosus ihienvmcwni Koenike (S/alays 
1932), or in Eutromhidium otorheiense Feider (Feider, 
1946k Simple shortening of a leg, but without othci 
abnormality, has been recorded by Feider (1946) in /:. 
otorheiense* and termed micromely. Schi/oniely 
(complete or incomplete duplication of a leg) has been 
recorded by Andre (1949, 1960b) in the trombidioid 
miles Mkrolrott/bidium sucidum <L. Koch) and 
CQrpnrfirombium carduigerum (Berlese). Various leg 
and other abnormalities m Hydtaehncllac and other 
mites were recorded and discussed by Thor (1926) 

I * re of the developing eggs of other arachnids 
|0 supraopiima! icmperaimcs has produced limb, eve 
and other deformities. Among the limb deformities are 
braehymely and .schi/omcly (opilionids: Juhcrtfue, 
1968; spiders: Jacuriski. 1971, Mikulskaand Jacuriski, 
1971, Mikulska, 1973). 

The observations reported here indicate Thar 
developing limbs of Acarina are sensitive to teratogenic 
agents, as yet unknown. 

ClussiJ'icuTion of Post-larval TrombeUa 

K nown post-larval forms of Tromhetlu s. str. 
comprise six closely related species, characterized by 
Ihe lack of a crista metopiea and the presence o\' large 
idiosoma! depressions, allotted a glandular function bv 
scune authors. The genus and species were founded on 
a single specimen of T. glandidosa, found deeply buried 
in soil, Adria, northern Italy (Berlese, 1SS7). This 
specimen, which has noi been resiudied by anv later 
author, had several unusual morphological Icaturrs 
not recorded for any oilier species since placed in 
TwmbcUa s sir; (I) the presence of a chitinimi pore 
in each of the idiosomal pits, whether dorsal, lateral 
ot veiio;il, ( jphe dorsal idiosomal pits not contiguous, 

in fact well separated and occupying only a- small 
proportion of the surface; (3) the posteromost median 
dorsal pit being set in a group of four of the lateral 
pits, instead of being set m ihe midst of a group ot 
six pits; (4) the picsence of a long sinuous idiosomal 
seta arising from a small prominence on the lateral side 
of each propodosomat sensillum. These features ol 
Berlese 's original drawing were reproduced in Thor and 
Willmann (1947), and the sole further specimen of 7^ 
xlunditloso recorded, Austria, by Schuster (i960), 
is stated to correspond accurately in all moiphologieal 
features they give, apart from being somewhat smaller. 
One may therefore accept Berlese's drawings as 

The African T, favosa appears to be the species most 
separate from 71 glanchdosa. Dorsally it carries large 
idiosomal depressions, all lacking a central pore, bul 
surrounded by a prominent ring of ehitin. The dorsal 
idiosomal setae are also markedly different, being in 
the form ot large flattened scales, the scobillum ending 
in a long central point and being provided with iarec 
projections along its lateral edges (Andre 1936a, 1958). 
In T. gluhdtdosu the dorsal idiosomal setae arc 
spinelike, simple, and bent near ihe base of the shaft 
so that the main part of the shaft is parallel to the 
surface. In 7? lusi/anica the general idiosomal setae arc 
spiiiiiorin, simple, curved, sharp, on short tubercles 
(papillae), only the more peripheral ones of the 
propodosoma bearing fine setules. lu T. warregensis 
the dorsal idiosomal scia is papillate, elongate, eui~ved 
near its base, and ornamented with prominent broad 
setules in (he form ol scales or long barbs; in both T 
adelaideae and T. alpha the seta presents as a tear-drop 
perched transversely upon a papilla. In 77 adelatdeac 
the scobillum is ornamented with prominent broad 
scales, while in T. alpha only a reticular pattern is 
visible (see Figs 7B, 9 AD) 

The other hitherto recorded species of adults of 
Tromhd/a T, lusitanica Andre. T. warregewslg Hirst 
(which appear to be close to each other) and /. 
adelaideae Worn. — differ in having dilfcrcul members 
of the idiosomal pits with central pores, in setal and 
leg characters, etc. (One feature used by Andre in 
proposing T. lusitanica and separating it from T. 
ttlandtdosa and F warrezensis, was stated to be its 
possession of a punctate seta-less area on the 
propodosoma. However in 7^ warregetnis re- 
examination of the holotype under oil immersion 
shows minute rjunctia in this glabrous area. The same 
i iImohs area [tl T, adelaideae and T, alpha has small 
rounded tubercles.) 

Thus in T. adelaideae nn<.\ T. alpha the dorsal pirs 
caTtv ihese ehiriuoiis pores or rings more or less 
centrally in lateral depressions 2, 4 and 5; in T. 
wamyensis thev arc present in lateral depressions 2, 
3, 4. *i and b; m A lusifutiaa ilicy aic picsent in lateral 
depressions 2, 1, 4 and 5 (Andre, 1944) 






l [< i. y. DOdrs&l idiosoma setae in adult Tromhella. A, B Trombetla adelaideae Worn., Holotype, A setae from the anterior median dorsal 
depression; B setae from the posterolateral edge of the dorsum. C,DI warregensis Hirst, Holotype. C Group of setae from posterolaleral 
part of dorsal idiosomu; D Two setae in lateral view at the posterolateral edge of the dorsum. (All to scale shown.) 

The arrangement of the setae of the dorsal idiosomal 
pits has been used by authors (e.g. Womersley, 1954b) 
as taxonomie criteria. Thus the central areas of these 
pits in T. favosa bear slender papilla-less spiniform 
setae (Andre, 1936a), while in T. lusitanica the pits bear 
spiniform, conical setae arising from a short tubercle 
(Andre, 1944); in T. glandulosa they are short, conical 
and curved (figured by Berlese, 1887). In the three 
Australian species recorded here the setae of the pits 
do not differ essentially from the general dorsal 
idiosomal setae. In both T. adelaideae and 71 alpha the 
peripheral setae of the dorsal pits may form a single 
vague row, but in T. warregensis the central part of the 
dorsal depressions is nude, with peripherally the setae 
in one or two fairly regular concentric rows (in T. 
g/andidosa Berlese figures one row of setae near the 
centre, and one row at the rim). 

Some use has been made of the relative shapes of 
leg segments and the rations of various segmental 
lengths as taxonomie characters of trombidioid and 
other mites, although with soft-bodied mites such as 
the trombidioids too much reliance should not be 
placed upon the shape characteristics of the tarsi. For 
the three Australian species of Trombetla the ratios of 
tarsus 1 length/tibia I length and tarsus 1 length/tarsus 
I width, are given in Table 6. 

The data in Table 6 indicate that T. adelaideae and 
T. alpha resemble each other more than either 
resembles T. warregensis, which agrees with the 
idiosomal seta characteristics. 

The data presented or previously published appear 
adequate to separate species of Trombella on a world 
basis, even though we do not have any species recorded 


RfcC S AUSr MUS 10(11): M5-LM 

.tow, MM 


T, WARREGLSSIS \%$* aWo T_.hi, 5 hoi RfeUic* to lengths) 


A< B712C 

alp hi 

i nvmplis* 
AC 1*7)20 


T, ack'laidi'Chl 

7i iwfcwjBff/i&fcj 

Tarsus ! 
Fihia 1 

l.irs;il width 

length Tat 

Width Tal 






1 59 



1 5(9 

lUKtmd >< Uttfffltft 

Hi <R> 

2S(t :h: 

100 102 

2.K0; ? 76: 


1 r f„,nUl ). ICVtytnik* 

11 I (Rj 

|7H 164 

:.::<:, 2x325 

* omitting hrachymeWc specimen AOtntZA- see tfaia m T.ibi<- 2, 

t Re-measured from hololype. Sec data in TaHle 2. In both leg I is seen in lateral view. 

3 Womersley (IV54b, pp. 127, I2K) etves these Itgurcs as 270 afld W) and the ratio ah 3 I I rtt M'tvnneii lias mot hern remounted. (In 

1939, p. 149, heyave these lignres as 260 ;nul 90 ) 
% Womersley < 1954b. pp. 126, 1281 (who bad remounted the specimen! (gVptLkcsfi figure* S3 JTJttld 155, and the ratio as 2.75:1. Hirst 

(1929. p. 170) gave 370 and 140, i.e. a ratio of 2.64:1. 

in which both adults and dcutonymphs have been 
described. The following key is therefore presented. 

STAG! s oi TuoMBEU A $ $TH 
I. All dorsal pits of idiosoma lack a central chitmi2cd pore- 
structure. Dorsal tdiosomal setae flattened, with long 
median end and 3-5 strong lateral projection.";. Central 
areas of dorsal pits with long spiniform setae, not 

mounted on papillae T. favosa Andre 

A! least some of the dorsal pits oi 1 tic idiosoma with 
a ceniral ehituu/ed pore-structure 2 

2(1) All dorsal tdiosomal pits with a central chitini/ed pore- 
structure. Dorsal idiosomal setae spiniform, bent near 
base so thai the shaft lies parallel to body surface. Ratio 

of length tarMts I /lib IB I ubuut 2,0 . . . . , 

i , — ■ , ... ... T, ftitwdtiimxi Rcrlccc 

At least the antcromosi of the lateral row of dorsal 
idiosomal pits lacking a central ehitinized porc-strikime 

3(2) Dorsal idiosomal setae long and pointed, the seta 
scobillum not wider than its basal papilla. Third lateral 
dorsal depression with central pore-structure ..... 4 
Dorsal idiosomal setae tear-drop shaped, the scobillum 
wider than the basal papilla. Third lateral dorsal 
depression without central pore-structure 5 

4(3) Dorsal idiosomal setae with numerous prominent 
setules. Chitmi/cd pores present m lateral row pits of 
dorsum numbers 2, 3, 4, 5 and 6. Tarsus l/libia I aboul 

1.2. . . - »• i * T warregensis Hirsl 

Dorsal idiosomal setae either spiniform or with slender 
setules. CiumiLzed pores present to lateral row pits of 
dorsum numbers 2. 3, 4 and 5. Tarsus l/tibia I about 
1.35 1 , . . . i ..... r. lusitanica Andre 

Vi) Scobillum of dorsal idiosomal setae with distinct ciliate- 

barbed surface 77 adrlaidcae Wornrrslry 

Scobillum of dorsal idiosomal setae with smooth 
surface, not presenting as a eiliatc-barbed structure 
. >, T. alpha n sp (deutonymphf 

Leg Chaetoiuxy of Trombclla alpha Larva 

Specialised sensory setae upon the legs of larval 
prostigmatic mites have been of interest to taxonomists 
for many years, particularly in classification. Although 
they function as physical and chemical receptors, then- 
have been few functional studies upon them, efforts 
having been mainly in description and class! Hcatum, 
with their functions only rarely being postulated. 
Classification of these setae rus been contentious, with 

two main systems being used in the terrestrial 
Patasilengona, which I have designated the 
"itombieulid system" and the "Grandjean system'* 
(Southcott, 1961a). Despite these differing class) ficutory 
systems, there are no doubts that seial patterns differ, 
not only betWOfifl families, but between genera. Within 
the- L'enera allotted Lo the family Trombellidae 
considerable differences in leg setational patterns occur 
(Soi.ihcott, 1982). 

Although the presence of modified "normal setae" 
or scobalae — 'taasLkette" — is well-known on the 
femora of trombiculid mites (Audy, 1954; Southcott, 
I96ib)i the presence of simple sensory setae upon the 
femora is more restricted. In the TYombellidae, in 
addition to being present on femur I, II and 111 of 
Trombclla alpha, they were recorded earlier on the 
telofernora I and 111 in Raiphaudyna (as nude 
femoralae) by Vercammen-Grandjean et al. <1974). 
However, they occur more widely, being also present 
upon the femur of larval Ncottumhirfium (Southcott 
1954; Borland, 1956) and Mutwitguis (I .indcpnsi un<\ 
Vercammen-Grandjean, 1971) (Ncotrombidiidae or 
Neotrombidnnae), in Durenia (Vcrcanunen-Grandjean 
1955; Vercammen-Grandjean and Audy, 1959) (TVom- 
bellidae) and Hannemania (Hyland, 1956) (leeuwciv 
hoekiidae or Lceuwenhoekiinae) and more widely in 
the Trombidioidea, such as in Megophthromhium 
(Mullen and Vercammen-Grandjean, 1978) (Micro- 
trombidiinae or Microtrombidiidae), and in various 
genera of the Johnstonianidae (Newell, 1957; Robaux 
I97K), It would thus be unwise to place too much stress 
on the presence of these setae on the femora of the 
ktrval instars as a taxonomic character. 

Similar smooth sensory setae occur also on the 
adults and nymphs or Ihjmhetla (lug. 8A-T). They also 
occur more widely in the Prostigmaja eg Smarts 
conpen Southcott (Smarididae), referred to as spinulac, 
spinofernoralac, etc. (Southcott, l%lb), and on genua 
I-IJI of the nymph of Mierotrombidium fflrsutUfti 
Worn. (Southcott, 1946b), 

A not her resemblance between Trotnhetla and 
Raiphaudyna ^ larvae lies in each possessing two palpal 
femoralae. In Tromhella, however, there are no 



vestigialae; in Raiphaudyna these arc present as distinct 
'^mushroom-like mtcrosputV (Vereammcn-Grandjcan, 
vi Qt» 1974. pp. 248-9). 

Generic Ci&wtffeMofi of the Tr&mhtHtdite 

1. Adults and nyruphs 

l hor (1935) defined the Trombellinae as Those 
Trornbidiidac which lacked a crista, with eyes 2 | 

2. sessile, setae shurl and sharp, wifh two sensoty setae 
on Ihe propodosorna at the level of the eyes, and some 
lesser characters, ro include only Trombelta Berlese. In 
1937 Wotnersley added Chyzeria Cancstrini 1897 to be 
subfamily, as well as Puruchyzeriu HirsI 1926 (syn. 
Thuamaloihrombium Andre 1938") (1938b)); this was 
accepted by Thot and Willmann (1947). However, both 
Parachyzeria tndira Hirst and P, poecilotrichum 
(Andre) have two pairs of prosomal sensilla, and 
therefore should be placed in the family Johnstotuan- 
idae Thor 1935, subfamily Johnsronianinac as defined 
by Newell (1957). Womersley (1954b) described the 
reared eyeless dcutonymphs of Audyana Worn., 
dividing the subfamily into the tribe Chyzeriini for 
Chyzeria and Parachyzeria, and lVombellini for 
Trombella t Audyana and Nothrotromhidium Worn., 
1954 (erected for the Furopean T. utiontm BerL), but 
did not redefine the subfamily. Vereammen-Grandjean 
(1955) described the reared nymphs oi \' Durenw, and 
provided a revised key for the tribes and genera of the 
post-larval Tiombellinae, Feider (1955) elevated the 
Trombellinae to the family Trombellidac, a status which 
most later authors have accepted (see Southcott 1982). 
Within this family (or subfamily) two further genera 
have been proposed for adults or adults and nymphs): 
Parathrombella Andre, I9SK and Neonnthrothmm- 
hidiam Robaux, 1968. 

In defining these two last-named genera each author 
staled thai a crista metopica is present. In 
Parathrombella nasuta Andre, the type .species Of Us 
genus, the propodosorna bears dorsally I i I eyes 
laterally, and in the central part there is a thickened 
ehitinous plaque, roughly in the shape of a trapezium, 
with prominent anterolateral angles, each bearing a 
sensillary seta, which are thus well separated I here is 
no anterior linear projection of chitin from this plaque 
oi shield- The hyslcrosoiua bears KMor i I) contiguous 
plntes, which resemble (hose of Tramhellu, there being 
lout 111 each lateral row and two or three median. Andre' 
therefore ledefined the Trombellinae, saying that the 
crista was limited to a more or less developed transverse 
band, with v 'rarcmcnt un vestige, trcs re'duit, median". 
The use oi the term crista etc. seems to be largely a 
matter ptf terminology. I here te no doubt that the two 
species Andre classified in Parathrombella belong to 
the Trombellidac. However The other two species thai 
In- pl.u ed In Parathrombella differ from P nasuia in 
having the following three characters: <1» eyes 2+2. 
(2) no (arge dorsal hysterosornal depressions but instead 
a hexagonal network connecting the bases oi the seta 

papillae, (3) two small circular pits amerolaleralfy on 
dorsum of hystcrosomu. Andre (1962) recognised that 
the two latter species come within Durenia, 

Robaux (1968) erected Neonothrothrombtdmnh WWl 
type species N. Jranzi Robaux, 1968 from South 
America, stating that in this genus the ensta is well 
developed, but did not discuss any consequent revision 
of the definition of Ihe Trombellinae As earlier (I%6) 
he had referred to a crista in Nothrotromhidium 
otiorum (Bed.) as consisting of no more than The two 
prosomal sensillary bosses, he was clearly using the 
term crista ffl a wider sense than the majority of 
authors. Even so, he later (1968) figured in N. jranzi 
a broad ehitinous thickening of the dorsum of the 
propodosorna extending from its rear border, anterioily 
enclosing Ihe sensillary bosses and with an anterior 
gutter-like seta-less delimited extension running into the 
nasus. In the second species he placed in this genu*. 
N. americanum Robaioc, 1968, the propodosorna bears 
dorsally an oblong plaque, extending only as far 
forwards as to enclose the sensillary bosses (which are 
anterior to the eyes), and without any further antei ior 

In my opinion these large thickened areas on ihe 
dorsum of the propodosorna correspond to the dorsal 
shields of other prostigrnatic mites, and could well be 
termed a scutum rather than a crista. The same 
situation occurs in Chyzeria, where there is evidence 
of a dorsal propodosomal scutum and also of a 
rudimentary crista (Southcott, 1982). The crista may 
be considered as absent in N. americanum, and 
rudimentary in N. frart;.i, 

The differences enumerated between N. /rami and 
N. americanum justify their generic separation, ,m;i 
for N. americanum the genus Maiputromhetla n. gen. 
is proposed. 

Although the definition of the Trombellinae given 
by Thor (1935) and Thor and Willmann (1947) included 
2 t 2 eyes, yet eyes are absent in Audyana, present 
as I ♦ I in Parathrombella, and as 2 ^ 2 in Trombclla 
and other genera. For Nothrotromhidium Worn, the 
status of (he eyes is unclear, as Robaux (1966, Fig. IDj 
shows only 1 » I for N. otiorum. the type species (hut 
does not comment on this in the text), while Andre 
(1960a) records that N. hreiuarsum Andre, I960, has 
2 4 2 eyes. 

From the foregoing, a revised definition of the 
Trombellidac may be offered, with also subfamilies, 
and a definition of a new genus. 

Family Trombellidac Thor 
(Originally as Irombellinae Thor, 1935) 

l>c fi n 1 1 i n n : Tro m b l d i o i d c a with absent m 
rudimentary crista metopica Fyes absent, or present 
as J 4 I or 2 r 2; if present, sessile. Propodosorna 
dorsally with a pair of sensilla (trichobothria), Sti 
a fossa which may have an elevated rim. or be present 


REC. S. -MJST, Mli& Will: I4S.I&H 

JW. /ftfo 

as a papilla wit b a central hole. Dorsum Of 
propodosoma commonly with a thickened scutum, 
which may be poorly defined at its borders. Idiosoma 
commonly highly modified* with pits or depressions 
with a possible glandular function, and bearing 
modified setae. Body and leg scobalae borne singly, or 
in groups upon small cttticular plates. 
Type genus Trombe/h Berlcsc, 1887. 


J. Dorsum of hysterosoma with prominent seta-bearing 

projection* . ...» Clo/criluae, nev* status 

Sole genus - - . Chyzerio Canestrini, 1897 

Dorsum of hysterosoma without prominent seia-be.inng 
projections, but may bear depressions ot possible 
glandular function (Trombelliuae, v str.) 2 

2(1) Dorsum of hysterosoma with pits or depressions , 3 
Dorsum of hysterosoma without pits or depressions? 

Ml) Dorsum oi hysterosoma with two small anterolateral 
pits. Propodosoma may bear a ehhinued plaque 
riot sally, adjoining or enclosing the propodosomal 
sen.silla. Eyes 2 -♦ 2, about level with sensilla . 
, . , . , .... Durenw Vercammen-Grandjean, 1955 
Dorsum ul hysieiosoma with three longitudinal rows of 
circular, oval oi oblong depressions , . . , 4 

4(3) Six depressions in lateral dorsal hysicrosomal row. Eyes 

Ja 2 Irombelkt Btrieve. IHS7 

Four depressions in lateral dorsal opisihosomat row. 
hves I t I - . Parathromhelta Andre, 1958 

5)2) Dorsal idioNoinal setae on papillae, in groups of 2-12 
in small cuticular plaques Sensilla well separated, Lyes 

absent Audyatw Womersley. 1954 

Dorsal tdiOvomal setae on papillae, arising individually 
from cuticle . - 6 

6(5) Dorsum of propodosoma Without significant thickening 
lo a scutum or plaque-like structure. Prupudostnnal 
sensilla close together, somewhat posterior lu level of 
eyes ihu N. otmrum (Berl), from Robaux. 1966) . . 

. . , NofhrotrombUJiutn Womersicy, 1954 

Dorsum of propodosoma with a well-developed scutum 
or ehidnizcd plaque in its postcriormedial pan Brts 2 

4 Z i • • i i ■ • • * • * * • t • < • * • ■ 

7(6) Propodosomal sensilla approximated, level with eyes, 
on an elongate propodosoma! scutum which extends 
tot ward on to nasus, there forming a median furrow 

Neonothrothrombidium Robaux, 1968 

Propodosomal sensilla well separated, anterior to level 
of eyes. Propodosoma bears dorsally a large transverse 
oblong scutum or pku|ue, enclosing sensillury bosses; 
there is no anterior extension of this scutum. Palpal genu 
bears rfistally on its lateral face a row oi broadened, 
spatulafc setae. ....... - Maiputrumbel/a u. gen. 

Definition of a New Genus oj TramheMdiw 

Mxiputromhclte n. gen. 

Definition; Trombellidae with 2 *■ 2 eyes, posterior 
to level of propodosomal sensilla. Propodosoma bears 
dorsally a large transverse oblong scutum or plaque, 
enclosing the sensillury bosses; anterior chitirwed 
extension of propodosomal scutum absent. Body setae 
short, blunt-ended, on short papillae- Palpal genu bears 
distally upon its lateral face a row of broadened, 
spatulate setae. 

Type species Neonothmihrombidium amerieafwrn 
Robaux, 1968. 

This genus is known only from its type species. I lie 
generic name is derived from Muipu, Chile, the place 
oi' collection, and Tromhella. 

(2) Larvae 

Southcott (1982) discussed the characters of larval 
Tmmbellidae, concluding that it was not possible to 
give a formal definition of a larval rrombellid Of ibe 
two subfamilies into which the Irombellidae is here 
divided, the reared larvae, except Chyzeria, belong lo 
the Tromhellinae, 

A review ot the characters ol the larval fromhellidae 
shows thai Womersleyia Radford has a number of 
features in common with Durenia, so thai I have 
decided to restore the former to full genetic status (see 
key below.) 

The following is a revised key to the larvae at present 
allotted to Trombellidae. (Raiphuudynu Vercammen- 
Clrandjcan et ul. % 1974, has been included, although fl 
review o( its characters, with two pairs of scutal sensilla, 
shows that its affinities lie with the family Jolinstonia 
mdne. near the subfamily Lavsemiuae Newell, 1957, 
Where it is provisionally placed i 


I I eg segmental formula 7, 7, 7 . 1 

Leg segmental formula 7, 6, 6 Or f ». 6 ( h . ,4 

l(\) Dorsal scuium lacking anteromedian proiccli'Mi 
("nasus"). . Chyzerw Canestrini, 1897 

Dorsal scutum with anteromedian projection ( ,v nasu03 

\{2) Dorsal scutum with 8 seiac 

, . , kutt>huudytia Vercammcn-Grandjean 0/ at, y 1974 

Dorsal scuium wilh 6 setae ( , 

Noihotromhicula Dumbleiuii, 1947 

4(1) Leg segmental formula 7, 6, 6. AM and AL (latter close 
to Sens) setae short, elavate. Two claws to each 
pedniar.sus Palpal tibial claw with lour prones 

| Vctecunnicn-Grandjean, 1972) 

i Audyunu Wouicrsley. |S54 U lJ >4a» 

Leg segtneuial forniula 6, 6. 6 . - - . . .5 

5(4) Tarsus ot each leg with a single claw , . 6 

Tarsus oi' legs I and II with g single apieally trifutcatc 
claw, taisus oi leg in with two claws . - 7 

6(5) Scutal geftsiUa well hrhind level ot AL \cutabt\ about 
midway between levels oi' AL and PI . Cheheerac 
compact, the combined chelae bases not more than twice 
&S long as wide ... . . , - TmmbeUa Bcrlcse, 1887 
Smtal sensilla only a little behind level ot AL scutalac. 
C'hdioera* elongate, the combined chelae bases more 
than twice as Uhhj as wide (for N. otiontni (Hert.j. trom 

Lender, 1958) 

,\uthro{rombtdiutn Womersley, 1954 (19Mb) 

7(5) NaMis ot -.euiuni small, laigely occupied by itlC ba^es 
of (he AM sculalac, auJ with a deep constriction hehind 

l,eg tibia 111 with a large solcuoidala 

. . . Woniersteyw Radford, 1946 

Nasu^ al seitium lat^e. Itian^ular, its lateral borders 
eoniinuous wilh anterolateral borders of scutum, with 
at most only minor constriction Leg iihia 111 without 

a large sotenolttala 

Ditn'ntu Verca.iuiicii I .kuilIjcuu. [$53. 



fa) Function of Tarsi in Irombellct alplia Larva 
The attenuated latsi of larval T>ombeila alpha 
represent an unusual shape among lai val prosttgmatie 
miles, suggesting that I hey mighl serve as sound- 
receptor organs. If so, they could possibly be usetul 
in locating males of the hosi cricket. At my request, 
Mr Davidson, the collector of the specimens and data, 
has provided information on the numbers of mites 
parasitic upon male and females of Teleagryllus 
commodus, as well as for sub-adults (At and A2 
insiais). As only the adult males stridulate, for the 
hypothesis to be valid there should be a greater 
tendency lor parasitizatiou of ihe adult males. In Table 
7 data are provided on the numbers of miles on the 
various instars of the crickets, by date of eollection. 
r:\itmiuatiou ol the data in Table 7 shows that there 
is a greater number of larval mites attached to the adult 
male nickels than to aduli females, the proportion 
being 3.70/1.15 overall, or greater than 3:1 Submiitmg 
the data to a test of the null hypothesis, i.e. (hat the 
proportions of the mites on the crickets are 
independent ol sex, we find that X 2 - 83.52 on 
I d. f., i.e., P<.0<)1*** 

A significantly greater degree of parasiti/ation of the 
adult male criekets occurs than in the females. It 
should be pointed oui also that as the crickets were 
captured in pitfall traps (using Vaeola jars as rrapsi. 
there could have been some transfers of mites between 
the crickets, since these traps functioned over several 
days and the conditions in the traps the crickets were 
extremely crowded when many were caught. Although 
only a proportion of the mites were studied, among 
the samples suhmincd there was only this one species 

oi larva. 

I his ti riding therefore is consistent with the 
hypothesis that the miles may find their hosts, at least 
in part, by the use of a sound-detection mechanism. 

Since only the adult male crickets stridulate, the 
hypothesis may be examined further by comparing the 
figures o\' parasiti/ation for the sub-adult criekets; the 
firSI and second insmr sub-adults are the only sub-adult 

insiars in which the sex is readily determinable. (Mr 
Davidson advises (pcrs. comm., 1983) that difficulties 
in sexing account for some differences between the 
totals in the tables.) 

Table 7 shows that in the sub-adult instars the mites 
appear to have a preference for the female crickets, wuli 
a mean of 1.57 mites per female against a mean of 1.25 
mites per male. If however we compare these figures 
by the same null hypothesis as above, we calculate that 
X 2 = 2.90 on 1 d. f., not significant, i.e. the null 
hypothesis is not disproved. 

There is thus no evidence o\' a sexual pteference by 
the mites m parasitizing the sub-adult crickets. 

These results thus are consistent with the hypothesis 
that the mites make some use of sound-detection in 
their searches for their cricket hosts. 

fb) Possible Auditory Organ* m Acarina 

It has been suggested above that the larvae of 
Tromhdla alpha use Ihe leg tarsi as sound-receptor 
organs for locating the sound-emitting adult males ol 
the enckers, winch is supported by the greatei incidence 
of parasitization in adult males than in adult females. 
Other instars of the crickets are also utilized by the 
miles (compare Tables 3, 7). 

Earlier suggestions have been made that there are 
structural adaptations in ectoparasitic mites which 
serve an auditory function in host location. Newell and 
Vercammen-Grandjean (1964) described two species of 
miles from Africa in the family Johnstonianidac 
(Trombidioidea) as Pteridopus auditor Newell and 
Vercammen-Grandjean, 1964, and P, pseudohatme- 
mama Newell and Vercammen-Grandjean, 1964. fl 
auditor had been collected parasitic upon cricket^ 
while P. pseudohannemania was described from a 
single specimen collected free on the forest floor, 

In both ol these species there is an unusual 
morphological arrangement and modification of setae: 
tarsus III and tibia III carry a dorsal row of long 
feathered setae, each o\' which has a preformed fracture 
line near its base, corresponding to an interruption of 
the aciinoehitin. They proposed that these setae 

iaiu I 7 

(Only live crickets examined) 


( ai'juorv 

\du1i Males 

Adult Rma 
No of No. ot 


Sub-Aduli* Ma 


Sub Adult* Females 

DstU "f 

So, ol 

No. oi 

NO; Ol 

No. nl 


1 fKlsCK 


\m N.. 






Nin Nc 



\..i V 







N p 












' 1 ■■ 






7 T 75 







2t.ii 79 


















— 7M 










14 U 7m 














































| ■, BO 
























Vluli I iintl mliilt 2 iir-uuv 


RBC S. AUST. MUS I9<il> 145-lfcS 

,&*& i$«W 

subserve an auditory Junction. aJlowsne :he larvae 10 
locate their hosts bv a sonar technique. These setae are 
generally broken off in the older (ie. fed) larvae. These 
authors proposed rhat Oris row of setae served no 
further useful function after attachment to (he host, 
(This would be consistent with the general finding that 
such larvae feed fully on a single host.) Additionally, 
rhe tarsi of t* pseudohunnewaiitu (hm only tarsus III 
in R auditor) are highly attenuated, each tarsus ending 
in a short pedk-ellus and three claws. 

The arrangement of the setae was considered 
suggestive "of the setae of rhe antennae of male 
Culkidac, which are known auditory organs". 

In a later paper Vcrcammen-Grandjean et a/, (1965) 
discussed another form of setal modification in the 
North American lecuwenhoekiid mite, Whanoma 
gtenni Brcnnan, 1962, which is an ectoparasite of two 
species of bats of two families: Phyilostomatidae and 
Eraballuronidae. In this species of mite rhcrc are greatly 
elongated setae C t mastisetae ,f ) on the larval tarsus III. 
These anisotropic setae, identified aa toys' itaTsalae, 
have two conspicuous bends and points of weakness. 
Most specimens of setae Inom mites taken in the field 
arc broken (92.1% of J4I setae), a situation analogous 
to that seen in the genus Pferidopus, These authors 
speculate that such setae assist in locating sound- 
emitting vertebrates, such as bats which emit their high- 
pitched sounds even when roosting, when almost all 
of the opportunities of attachment of these mites can 
be conceived to occur. 

Although these authors do not mention it, some 
other prostigmatic mites have highly modified sciac 
which might conceivably serve as sound-detecting 
organs. Thus in the family Erythracidae the genera 
Fxitomorto Cambridge, 1898 and Putophus Berlese, 
1916 have such upon the tibia IV of the adults (and 
possibly also on genu IV), With regard to the possible 
function of these setae, there termed "plumalae" the 
author (Souihcott, 1961a, p. 48ft) accepted the views 
of previous students that these setae were used as sails 
as a means of progression, fr is now apparent that a 
more detailed study of their function is warranted. 

Tn the case of Trombetto alpha larvae, the setational 
pattern does not suggest that the setae have any 
possible specialized auditory function. However, I he 
attenuation of all tarsi does suggest that they could 
function as sound receptor organs in host detection 
Such a function might be enhanced by the possession 
of a single tarsal claw to each leg. The use by this larva 
of a cricket as its host species is possibly of significance. 
Since many animals emit sounds \\k a variety of 
purposes, such as territory-establishment, echo- 
location of surroundings and prey, sexual attraction 
and other fund ions, it would appear thai the 
development of host-locating organs by t heir potential 
parasites is worthy of a wider study. 


I am indebted to Mr Steve 3, Davidson, Zoology 
Department, University of New Cngland, Armidale, 
New South Wales, for collecting and forwarding both 
preserved and live larval Times and data for the present 
study. The South Australian Museum has made 
available the holotypesof Tmmhella warrvgensts Hirst 
and T. udelaideae Worn ... and a paratype of 
Womersleyia minuta Radford, for study. I thank Mi 
L. C. Veitch, Principal Research Scientist, CSIRO 
Division of Mathematics and Statistics, Glen Osmond, 
South Australia, for aid and advice in statistical 


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|V ufhrombtdiam ndnrhwrise I cider Amtt Acad. Rom \!r>n 

-SW|. Stitnl (3) IV. 1-44. 



i [ LOi K. /. ''. ,: *i i ;iia':i IVomtathitac nni p iwx to Iratf, fr. 
/? &>m v Ml. 

I t mi R. / 19*5 Vcanna Iron.bidotdea htiwa U. f Rom. 5: I 186 
midi !■:, /. |958 Ptipifi larvfi iiin ftunvlta irombollidni? (Acanna) 

tlbUlliilS pun jiliinJ M diHpre car icic. i/are a lainni.-i 
titfrf, fi ft ft /■//. toy* Start :n Cw_ Jftft ftirt >' H Affrti. 
'>: J65 2S2. 

I I tULk. Z. 1559a, Liujc dej earaertr^s *pniwh vtew Ics 

irombidioidea. AraralGg'tu J: $6£5. 

» HOHR, /. 1959b. (New proposal* on the .la-.tlic-uioi. Of ntiics 
from the BTUup Iiom' l idiai. ■■'"<*/, /hum 39 5J7-54S (l<> 
Russian, moimnry in r^nglish). 

MAUBNKBI R C* v. Ipm/^QgMieMwaetten. h-rti. z**o(,-tn>t. 
i.ey. Wen 18 H8M0J 

unit ; \i I ., I oomis, k. n, wr LBfcURN, w. C, and WRI -J 

W .1. 1982. A glov.uiy u| CfalgUtfT IftfhlOOiQlQi '• A " 
IVoinbicululaei, ./, /ncil htttotnof. 19; 22I-23N. 
HI NK.ING, H- 1SS2 DeJtTtlgC - CUl Auriioime, I t.PvicKUipgsgeM.lnchiL 
und Hiolofie von Imttthif/mtti HWvntuwitn Kent] /- hmv Am/, 

37: ^VdM 

H1KSL s. tW6 D«&tjplA>nt <>t n<u h.m.s, inctu4ifl__ r<»ui &ew RflBcics 
oi "red spider". Pwc />'"/- SOC London. l**26: K35-S*4| 

HIRST, S. 1929. Additional note, on Australian rmtKS el the latmlv 
Tiombidudnc. wlili Llffbcrlpnortfi oi new forms, fbfd, 1920: 

165 176. 
HOT I I I. INC., H. TO?, AtnK^* >»! .cmiiplev pi stcpshcal w.naOles 
into principal component. ./, vt/tn; Ps^\1u>t. 24- 417-441; 24 


rlYLAKU K. h. Ii; 1956 A new sp.-Ues ol ellipse/ mite. 

fUmi\,ituinut ficv^m-ri (Acaiin.t. Ir hiculi'Jaei. 7 l\na\i(t>l. 

42; 176 179, 
IA<. INSKI. 1 . 19?!, .ernpcraluie induced developmental 

monstrosities in /Iw/W/TO alr/Cfl ( - I Koch tAraneac, 

AgjClClfUkcJ. /_»/. flpi 21 :M:-V7 
lOHNsrON. I) P.. and W.Vkl.R, R H. 1967. Observations on 

po-acnitn viipiL dewlupmciu m EUtrrtffihiCUfa spfrffrftaf (.Vari- 

v.op'VTnc*.) / "«■</, ( nt&mQl A- JOG^ID- 
H iM.s, U. m 1951 I he growth of Itie harvesa nnic. rr<<mbiattu 

uiititti/thifts siia'.'.- ftyrvsltatirgy 41 229-248, 
JONLS. 1*. M 1954. Op ilv rote "i" the iMQgunicni III acJiinc 

dcvoloptncni and Hs hcaiinf on piip;tinrni:iiuin QlMff -^ (WWh 

\w. 95: H&I&J. 
Jl BhKTHIt, C I9ft8 lrniliH.K'ic LApeiancnralf J.c- yti opdioti 

(.uachnidfh 7, Emttntal r\r- Marph, IV: 49*52. 
kran I /,. Q. vv. 1978. "A M.-rmai of Acnrtilogy n . 2nd edn< i htgpu 

Siaic Umvorsiiv BnpksiDnw, Inc.; Cot ".alii-,, Oregon, 
i INDQU16XHE: .'and \trc\m\u\ GRANfWtAW.IP Hi Wi 

Ko iMon ni nit clTi^tfi -lite larvae ol 'he aen^ra NeatnitnbhJiu+fl 

I fntKinli ;\n{\ Matmrtvjiis Wlinrltm, wit*i _i icdi ■I'mitUin ol the 

•aiblamtly Neoiiombidiiniit Kidct in 'he |Jontt>Hftid& ( V nm;r 
I'rusityiiLiia). (\uuta. Ftit(ftrif>i. IQJ 1557-159(1. 

MIKW1 SKA, I 1973 L-.peiifnt'piaUy induced Ucvclopmcnlal 
mOlWlm^iliw in tflC waiCr spider Ar? vrom'fit ^h-fuwia* H'^wku 
A>ot P&L 21 12h-O.V 

MIMJI SKA, I , :i...1 IAC'1 INSKI. I I97I. Ilu.hotonn in embryos 
oi lite --pidei Twnvria airka t I K.mJi JOdUfCd 'a 
sutuaopiipMi tcrnnenttutt LTeaiawnl iTi carl> «rfit»tyog$oy. A><9f. 

/W 21 21S1-2M 
MU1 I t N.i.. R ..and Vl:kr\MMFN-tiRANLULAN, r. M I6T78, 
\fi'i!t.>p/nhn>ttihiu/n. i HCT gtfHU3 in ilw I I) Ifombidudu 

(Ac;iu: I'anisiii-nyrma) "umi North Arncrioan MtoMpiiuH's. 

NI-AI I I and HARM I I . H < I9(d I In- l.lo rvcle of the ^rub 
ivphi^ chiwet nnic. ThQWbtruiU vkiUXWtil \tti fru So< 
Amvr. 54: m 203 

Nl WI! t , I \t 1441, Studios on Hie JutinMnr.MmJac lA..ari. 

Nl \M I I . I M , and I fcRCAMMEN-ORANQJEAN. P. H 1964 

PiefitititHS n j i Van. .lotm-.loraanidac) and |>rol>ahk* aildimi 

Or^MII in •■ mill-. Auitvttytiii *** ^H-MO 
RADtTi^D ( D I94h. laival and lYyiirphsl miles (Acanna 

liotnhu nliiJar) ffUin CeyfftH and (lie Maldive Island-- 

Putu;)t>'h>:v :.l 16ii4 
RtiHAUV, P. 1966. Sai iptekpirs M)roinbidiidac r.-ncs on FlOUveaUN 

i><iiM hi i.immc .u I ufiu- (\Ciri - 1 bn>mbidiidac). A^woln^to 

M Ml 63d 
RQBA1 'V I* »9r.7 i ..t.iihoH ,. lu.itli ie-, ... .iiicn-. 

I liioinhidii loi 'I'LuiLM't I. I tudt des ihiornbnJt'.n' jJulic--d«. 

!;i 1 Yiiiimili- llu'iiipic- II LiMC '.tuupiv* Efc<S lltri anhidi. -ns 

d'i iiroptf Mew Ktu* Vtt/i r/W/w rVtf?., rt//B f)i • I A [A»ol | 

4f> I J 2,, 
ROliAUX, P i'>ns rin ■.niL-idndjc d' du l 

liituiupodmac, MlllUPIIlanflfllft 1 liioiiibfllnn, i,V.:ii na 

Tliro'nhidiidiict. Act/hifo'titt ll> •1^(V-4hh 
ROKADX, H 1974 Rei hi ic NM air Ifi .JtVc|..ippcn.ciu « k blulCfjk 

:l" .it .men, "I hit .MthiUlnlaO ' M&Hl MUX, Mfirf '.''//''»/ *V(// . 

ftyftt, ffl s A A. AhjI HS 1-lKf.. 

koBAUX, ft I97H, (Jb^crvaiioiu ^nr tuidquBji Actinotftitt 

( Piostiymatcs> du sol d'Ameriqus du Notd. IX. Nbuvdlcs 

■ ijiwnif. dc trontbtdii-n-, ( Aiarij. \CtifQl(}f$Q 19: 2>S-J7| 
Si lUiSlFR,, R, 19C>0. AlliiLMicinr fan m,h ^\,j NjiLhncl.lLM, , ,.i 

Si-.-irnniv k (VllK \ririn-.pnda, Mid. mtlurw l*fr St6ktt' r W&h 

90: 5 7 
SOUTHCOn: R V, 1946a, On die Smandidav lAcanmi). 

frr>< Linn. '-»■ v.Mf: 70: I7M78. 
SOI HK.(.U l K V. I94f..b. Studies o^ Irornhidiidae (Acanna). 

Sv.ii.c oh.sctv.inon.s on the blClfeOgt ol the Mtciorromt'idHrun-, 

Pro, t >„n .Su.- \.sm: 7ft ^i:"-'k.. 
soi'TMcrrn, K. v I946C. Studies an Amtmliaji EtyrtnawJac 

(Acarlftfl)- f'nu. initt- Voj 1 - M£R 71 ft-4S 
soi.Mh *;M i. r. V W54. the fonus Neotramlitdium tAuaeitia: 

I ccuwcnhnckndae). L Ocscripnon ol tht ovtim and lana Hi 
\, ittr*'t»hidium humrivuno/i:^ fflrw I92M, tvllh an JCCOAm'l -a 
H.c bioloyv ..f H.r Lenii- fttfW & '*'" s 4 »/>./. 77 89.97 

N< fl IlKOn, K. V. 1955. The eflect ol .-"Id diocl. in pr.--duL.PK 
an.itcipl reratrrtoj^cal delwfniitcsineryihraeKJniTtw Twtu H 
5i/c. V a./.,/ 78- 169 i, r. v Nnia si.jdie--.Mi iWenytfcm&tt&aiulbtoItu) 

...I the t.tytlnaeoiden (, -A.rli j ci.a-'l rCVlUQT) id Uic 
cciuia and sublapuhcs. \i<sl. ./. /fool. \h 3C7-pip 

SOUIHCOII, K. V. 1961b. Deception Pi lwo pe\\ Au«tTOlinn 
s.naiid.dac iAcain...i, \uil. teniar*o. on cluteioiiiw and 
geographical distribution. Trans. Af. Sor. S. \t(sf K5: 133 I c - 1 . 

SOUIIKOI I, K V. 1963 I hcSrr.Hndi.1. i |fll North and 

< -. niral \tnciica and iowc oltttl counPi-> /rc/n. ft Sov, S 
4ii«, 80: 1^9 ^ 45. 

SOUTHCOn; k. v. iwu. feviwon or iin pnu3 CkttrfviontQ 

rmdein;nis (Acitr'tnrt: hi vl hraoidac). lUtf. >'- /"P/ 14 6JR-JMV 
SOL THCOTT, R. V. 1972. Mcmmoii of the larva, ol dte uiht 

C.Hlbdin.onKilini ( V.*ii.i[t: f.. wP.i.L-idac ) wiP' uhsL-u^tiotK on 

posl larval insist! iiiSt, J. cvO/i, Suppl. Sw '3 1-K4 
SOUTH* OTI, R. V, 1982. Obscf vaiKM.von t hvrxrui C ancspin. and 

some leMieti genera t vji ini rtombiJiotdefi) wiiii nenunki on 

rhe classiiM-UHin iP il.e -.aprrlainily ..nd ilescripiion id a 

pyyincphono mile photvp.- on Ctw:aui. ftee S". Aum \!u- leV 

s/,\I /NV", 1 , |933 ( \1issh.iduiu-' hffi WtU'l >hdr.uai an;. Ah// .4n v ;, 

'*'» 3)4-336, 
UloR, S. 1920. /-r/t/p M-mipnns ^. n. aus UracK^assct dfca Plm Ga-itsw aid, nebn eini^eii lienierkmiL'en ubtr 

Variabilitai unci MlisbUlhin^ bei den Acanna. /.mil AtlL *8: 


IIIOR, S. 19V-_ I ifvrstUil J ( "tMUilu.t^ del IjiiuIiC liOinhidlid.K 

W. I I cadi ISM m nnlcr hninlici. /<><>!. Ariz. W: L07-UX 
TFIOR. S., and WD t MWN, C. 1947. Irumbidndac. /Vv Tit>r,H,<h 

7lh. IS7-541, \xi\-\\'.vi 
VI-RCAMMRN-GRAND.IFAN, P H 1955. Un cenn: nouuan. 

Purcituj, dans la mu^famUI« da rrombcllinae iirotnbidiida.c— 

.Viii.iu: AV.' /not- Hot, off, bl 2""2-26(l 
Vbkc"AMMFN4"iRANDirAN,P U 1972 RevtMon dl W r | ( '\^;ii"i p (.* . | lvhwl-mI locajdje { ,ic I) Iii.mii the Asialic- 

Kieilic region. Folia porasitoL iPmhal |9 22 7 -25? 
\ I Rl AMMI N ckAM.-ir AN. \\ H 197.1 Sin les siaiuis dc In 

rami lie des TrambUlildae I each, IKI5 (AKarinn; Ptostigmawl, 
Aearotoria Ifi: 102-114 
VI RL AMMFN-CiRANrdr AN, f H.. and AtJDY. .1 R 1919 t!,w 
■veoiH_ir d i ^ e appartenant au genre Durvnia Vereantmcn 1955 
et originairedc Malaisie: Ihavma jHnttaporcnyfa n %p iVann,. 
Itouibid.idaei. Bialogweh J^arb. (Qpsdonaeal, Ghent 27: 98 Wl. 


1968 kevi-aon ot the Miblaniily Apolonnime W'liarmn. 1947 
(U-etj\venhoekndat v Acarina), Acomla^ia 1(1: 250-268, 
I M M KOIiAl'X, P, and SUZUKI, 0. 1574, tit\U>KmlyW 

it»iuttiLti\is, an nil .male homage let the mcrnorv ol I. Ralplt 

Audv tAcurina: Partiiiieiiffona) lap. / samt Zaai 25: 24^-249. 

VLI« \M\11 N (.RAMD.II AN ( P M., WATKINS, S. G., and BbCK, 
A J., 1965. Revision ot \\hanot\ui giCrVii 196: 511 
American bat para'.iic (Acanna: I ccuwcnhofkiidae). A*VtmlQ§fo 
7: 492 509. 

VI I /.I HUM, il. 1929. Ordain^ Mtlben, Aca.i In "Un: I .erwelt 
\1PteteoropaV , Vol. \ Section 7, pp E-LI2. <0uflil i Maym 
I eip/nil 

VITZTHUMi II ITOJ A. -in- fn "Hdndhuclt dei /oulopiC Llhc 
NaiuiecsclnclU-dcr Si U mmc vlcv i .crreUies" Ldv W KukentPal 
And 1. Kruinbach. Band I, Halite 2, I .eleruny U |>p. 1 I6U. 

VI 1/1 HUM, H. 1940-1941. Acanna. fn "KlassCTi iilid 0f4nUflfifn 
Jcs 1 icocivh-" lid M. <>- Uronn) Hand 5. Abicilun^ 4, fiech 
- (I Li._n.nn I 1 pp I 4Ml. 1940; J, pp 481-640. 1941; 5 6, pp, 
6-U-9).\ 1442 7, pp 9H-I0H. I04M 

WIL t MANN. v. 1919. Uic Arilitopodcrdauna von Madeira nach 
den l ijK-'.hin-M.n dtT Rcije voti Prof. Mr m. L.undblud Juh -Aukum 
L935. XIV. leireMnsche Acan (exkl Kodidae) Ark. f, /pp/. 


REC. S. AUST. MUS. 19(11): 145-168 

June, 1986 

WILLMANN, C. 1940. Neue Milben aus Hohlen der 
Balkenhaloinsel, gesammelt von Prof. Dr. K. Absolon, Briinn 
(I. Mitteilung). Zool. Anz. 129: 213-218. 

WILLMAN, C. 1941. Die Acari der Hohlen der Balkanhalbinsel 
(naeh dem Material der "Biospeologica balcanica"). Studien aus 
dem Gebiete der allgemeinen Karstforschung, der wissenschaft- 
lichen Hohlenkunde, der Eiszeitforschung und den 
Nachbargebieten. Biologische Serie No. 8, pp. 1-80 (K. Absolon: 

WOMERSLEY, H. 1934. A revision of the trombid and erythraeid 
mites of Australia with deseriptions of new genera and speeies. 
Rec. S. Aust, Mus. 5: 179-254. 

WOMERSLEY, H. 1937. A revision of the Australian Trombidiidae 
(Acarina). Rec. S. Aust. Mus. 6: 75-100. 

WOMERSLEY, H. 1939. Further notes on the Australian 
Trombidiidae with description of new species. Trans. R. Soc. 
S. Aust. 63: 149-166. 

WOMERSLEY, H. 1954a. New genera and species, apparently of 
Apoloniinae (Acarina, Leeuwenhoekiidae), from the Asiatic- 
Pacific region. Malaysian Parasites VII. Stud. Inst. Med. Res. 
Malaya No. 26: 108-119. 

WOMERSLEY, H. 1954b. On the subfamily Trombellinae Sig Thor 
1935 (Acarina, Trombidiidae) with the diagnosis of the nymph 
of Audyana thompsoni Womersley, 1954. Rec. S. Aust. Mus. 
11: 121-128. 

WOMERSLEY, H., and SOUTHCOTT, R. V. 1941. Notes on the 
Smarididae (Acarina) of Australia and New Zealand. Trans. R. 
Soc. S. Aust. 65: 61-78. 






The ovum, larva and adult of Odontacarus swani n. sp. are described, being the first recorded 
correlation between adults and larvae in Odontacarus. However, as is customary in this group, a 
larva has been selected as the type. 
A key is given for all species described as larvae from Australia, New Guinea and South East Asia. 




Honorary Research Associate, South Australian Museum, North Terrace, Adelaide, South Australia 5000 

(Manuscript accepted 23 January 19S5) 


SOI I I m OF!, k. V, 1986. Vhc genus a/otuueams lAcunnu: 
irombiailidae) II. Observations on the life history R0d 
morphology ol' QdottttHnruv swutU n. *p. and related forms*. 
Rev. S Ausi. Mu$. 19(12): 169-200. 

The ovum, larva and adult of Odontacarus swani 
n. sp. are described, being the first recorded correlation 
between adults and larvae in Odontacarus. However, 
as is customary in this group, a larva has been selected 
as the type. 

A key is given for all species described as larvae from 
Australia, New Guinea and South East Asia. 

A correlation analysis is made ol the dimensions ol 
the chilini/ed pans (scutum, setae and leg .segments) 
of the type scnes of rj. swani, A principal component 
analysis defined two components, both of which 
include variales derived from the dorsal scutum and 
the leg segments. 

A similar analysis is made for the larvae of two other 
species of Odontacarus: O. athertonensis (Womersley, 
1945) from north Queensland, and O. mccultocht 
(Womersley, 1944) recorded from north Queensland 
(and Papua New Guinea). In each of these species two 
principal components of correlation are defined which 
are not completely duplicated in the other two speciev 
All four of these components include variates based 
upon scutal dimensions, three of them include variates 
based on leg segmental lengths, and two of them (each 
of these latter being the second principal components) 
include variates based on lengths of idiosornal (body) 
setae. Some components contain also variates derived 
from body hair dimensions, 

It is concluded that no standard components can be 
defined and in general there is not a high degree of 
OOlteiatlen between the various standard (and other) 
variates customarily used in the classification of the 
larvae of the genus Odontacarus (family 
Ttombiculidae), in agreement with my earlier study of 
Trombella where another group of correlated variates 
was defined. This contrasts with similar analyses of 
hard structures (such as bone) of vertebrates. 

Additional morphological details are given for the 
Australian species of Odontacarus, O, cookt (South- 
cott, 1957), O. langani (Southcott, 1957), O, mathewi 
(Southcott, 1957), and O. southcotti (Womersley, 1944) 
to help construct the key of the larvae and for Ol 
barrinensis (Womersley, 1945). Amended collection 
data are provided for O. southcotti. 

A key is given for the adults of Odontacarus (all 
known adults arc Australian) and the adult O. snant 
is described. 

Acomatacarus retentu & (Banks, 1916), known from 
only the original material from Victoria, is made the 
type of Scopitrombium n. gen., in the Lecuwcnhoc- 
kiinae. A lectotype is designated, and the species 

Revised keys are given for the subfamilies of 
Trombiculidae, and for the genera of the subfamily 
Leetiwenhoekiinae for both adults and nymphs. 


The genus Odontacarus was founded by Ewing 
(1929b, pp. 22, 188) with the following definition: 

"Each chelicera with a row of backwardly directed 
teeth on the upper margin of chela, and some upturned 
teeth on the lower margin (Fig. 16). Palpal claw 
bifurcate, the two divisions unequal. Dorsal plate 
without median anterior process and with only five 
setae in addition to the pseudostigmatic organs, which 
are flagelliform and pectinate. 

Type: Trombicula dentata Ewing". 

Ewing's species was based upon twx> larvae: 
^Described from two specimens; one, the holotype, 
from Sonora, Texas (Bish. No. 10681), taken by O. G. 
Babcock, December 17, 1922 on a white-tailed deer and 
the other from Balboa, Panama on a cotton rat by 
L. H. Dunn". He figured (1925a, p. 258) only the right 
chelicera of the holotype (Cat. No. 888, U.S.N.M.). 

In founding Odontacarus Ewing <1929b s p. 188) 
compared it with the larvae then attributed to 
Trombicula Berlese 1905 (described originally from ihe 
adult) and to the larvae of Schongastia Oudemans 
1910, and saying "only the type species Included" i.e. 
T. dentata Ewing, 1925. He made no reference to the 
larva) genus Leeuwenhoekw Oudemans, 1911. 

In 1942 Ewing revised the taxonomy of some of the 
American mites Ccbiggers') then placed in the 
subfamily Trombiculinae Ewing, 1929 (1929b, p. 22). 
He (1942, p. 489) decided to divide the genus Leeuv>w- 
hoekia, and erected two new genera, Comatacarus and 
Acomatacarus. The latter he founded for Ihe larval A 
arizonensis Ewing, 1942, defining it as: 

**Most nearly related to ComatQcarus [Ewing, lac. 
at., p, 489] but differing from it as follows: Chelicera 
with a row of teeth on upper margin instead of a single 


RtC S. AUST; MUS. 19(12): 169-200 

Junc r 1986 

looth and both the tirsi [palpal femorala] and second 
[palpal gcnuala] palpal setae simple instead of one or 
both being plumose or barbed". 

The principal distinguishing characters ol Comoro- 
cwwy(from Leeuwenhoektu) were (Ewing, 1942, p. 489) 
"in having the posterolateral setae ol ilic dorsal plate 
of the usual form and similar to the anterolateral, 
instead of being elavate, and in having a large number 
of sessile setae on the dorsum of the abdomen, instead 
Of a small number situated on the tubercles". 

Ewing (1946, p. 436), in a further revision of the 
trombiculid mites (raised to the family Trombicuhdae 
by Ewing in 1944), redefined Odontacams and stated: 

"Odontacarus is related to Acotnutueurus Ewing, 
1942, from which it differs in having a single median 
seta on the dorsal plate instead of two submedian setae. 
It is also related to Endotrombicula Ewing, 1931 [See 
Wharton and Fuller (1952, p, 72} and Audy (1954, p. 
160) for opinions on the present status of this 
genus.]. , ,*; he included in Odontaco.nts also O, 
uusiralis (Ewing, 1929), which was described (1929a, 
p. 10) from several specimens collected on the lizard 
Th)pidunvz pentviamts at Verrugas Canon, Lima, Peru, 
15.jv.l92S, R. C, Shannon. Cat. No, 987, U.S.N.M. 

Brennan (1959) sraied that TYomhicuta dentata 
Fwing, 1925 was based on a single teratologies! 
specimen, possessing a single anteromedian scutal seta 
instead of the normal two, commenLing that this 
holoivpe specimen is in a distorted and damaged 
condition, with most diagnostic features hard to 
determine. Nevertheless he stated that 'The AM [setal 
is slightly to the right ot center", and declared that 
"there is no doubt whatsoever as to its generic status", 
even though its "specific identity ... is in question". 

With regard to the second larval specimen assigned 
by Ewing originally to T, dentota, Brennan (1959, p. 2) 
has commented that it is "damaged beyond recognition 
and reclaim .It seems incomprehensible, however, 
that this specimen could possibly have been regarded 
as con specific with the holotype". 

As far as O. austndis (Ewing) is concerned, Brennan 
(1959) redescribed the holotype briefly, without figures 
stating that this specimen also had a teratologic! 
dorsal scutum— "In each [of the two holotypesj, the 
left AM is lost; thus the specimens are freaks" Two 
other specimens, each labelled by Ewing as a "cotype". 
were identified by Brennan (foe. at) as Eutrambicula 
alfreddugesi (Oudcmans, 1910). 

However, eventually Q deniutus holotype was found 
not to be irratological alter all; Goff er ul (1972) were 
able to see two AM seta bases, using phase microscopy, 
although the setae were not attached (Goff and 
Loomis, 1977). 

Wharton -and fuller had earlier (1952, p. KM) 
rcftlfaod the probable synonymy pt' CkttMtUCttfilS and 
Acomufucarus, but had commented, with reference to 
Odonraeurus: "Specimens in existence ate too badly 

damaged to study satisfactorily" These remarks 
applied to both O. denfufus and O. australis. 

It appears strange that Ewing had not at this stage 
seen the characteristic feature of the leeuwcnhoekiinae 
shield, an anteromedian tongue-like projection to the 
dorsal scutum, in fact stating specifically that it was 
absent (see his definition of Odontacarus above). 
Although Oudcmans had described this feature in 
Leeuwenhoekta verduni (Oudcmans, 1910) as early as 
1910 and 1911, and again in his 1912 monograph, 
apparently it was not observed by Ewing and published 
until his 1942 paper for Avamatacurus and 
Comatacarus. However, in his key to the larval 
Trombieulinae, Ewing (1938, p. 291) showed that he was 
aware of the anteromedian process in Leeuwenhoektu 
Oudcmans, but again failed to record it tor Odorh 

Camaiucurus Ewing was reduced to subgeneric 
status (in Leeuwenhoekta) by Wharton el ul 0951), 
followed by Wharton and Fuller (1952, p. 96), Gould 
0956), Loomis (1956), and F'inlcy (1958). but restored 
to full generic status bv Reed (197.1) and Goff and 
Loomis (1974). 

Acomutucurus L.wing, 1942, was resurrected by 
Vercammen-Grandjean (1968, p. 122), and in its new 

i ric status was divided into subgenera by the same 
author— Acomataearus subgen., and Oroehlorus 
subgeo. (Vercammen-Grandjean, lot\ cit.). 

Odoniacarus has been divided inlo the subgenera 
Torsuiaratvs and Leogonitis by Vercammen-Grandjean 
(1968. pp. \2QA2\).Acomatacarus£w\n& 1942 includes 
several subgenera; Xenodontacarus Loomis and Got'f 
(1973) (a new name for AW/tfca/mGreenberg, 1951, on 
grounds o\ pre - occupation hy Xenaearus Kishida, 
1925), Mauritius Vereammen-Grandjcan, 1956 and 
Qrachlants Vercammen-Grandjean (1968, p. 122. The 
last has been synonymized with Tarsakaants by Golf 
and Loomis (1977), since the type species, 
Aiotnaiacams mkheneri Greenberg, 1952, actually 
belonged to Odontaairus* with the characters of 
Tarsaiacaruy which had page priority. 

The genera Austracarus Lawrence, 1949 and 
Hvruanus Lawrence, 1949, which were placed as 
si!'-y ( _'.._m ol' Aeo»nj><f<-<)i!ts hy Vcrcammen -GrojuhV.ui 
(1957), are now considered by the same author (1973a), 
without giving any explanation, to be foil genera m the 

With the restriction of the genus Odontuvurus and 
its separation from Atomalacarus, the only subgenus 
of immediate concern here is Leogoniu*, in which the 
Australia-New Guinea species considered here and O. 
audyi (Radford, 1946) are placed (see Vereamrncn- 
Grandjcan, 1968; Goff, 1979a, bL 

The genus Odontaauus as at present accepted is a 
widespread one, with species in North and South 
America, Asia, New Guinea and Australia. In the 
present paper the term will be used in the sense of Goff 
1 1979a, b). 



Most studies of Odontacarus species have been based 
on larvae, as in that mstar they are readily collectable 
as parasites upon vertebrates, including, in some 
instances, man. At times they are captured from a free- 
living stale, eg. upon cards, boots. Fence posts, etc. In 
a few Australasian species roarings have beer* achieved 
from larvae to nymphs. These were O. auUratlensis 
(Hirst), from Hollandia, former Dutch New Guinea 
(now Dyayapura, West Irian), reared by C. B. Philip, 
and 0. /ewgz/w (Womersley, 1945) (synonymized with 
O, novagutnea by Goff (1979a, p. 149)) from the 
Dobadura area of New Guinea, and O. nova-guinea 
(Worn., 1944), from "New Guinea", these last two 
having been reared by G. M Kohls. These rearings were 
recorded by Womersley (1945), and allowed him to 
define the post-larval characters of the genus (then 
Acomatacarus). Later larva to nymph rearings have 
been achieved by Dornrow (1956) for O. australiensis 
in Queensland, and by Nadchalram (1963) forO audy* 
(Radford, 1946) in Malaysia. The experimental rearings 
reported in Womersley (1945) allowed him to assign 
four species of adult trombidioid mites to the genus, 
these being O, attolus -(Banks, 1916) from Sydney, New 
South Wales, O. dromus (s. str.) (Womersley, 1939), 
from Long Gully, South Australia, O. pattim (Worn., 
1945), from Murray Bridge and Burra, South Australia, 
and O. retentus (Banks, 1916), from Lai Lai, Sea Lake 
and Ocean Grove in Victoria. Despite the widespread 
geographical distribution of the genus, and the fact I hat 
the larvae may be at times locally numerous, so far all 
adults that have been recognized as belonging to the 
genus are of Australian origin. Hitherto there has been 
no record of a correlation by rearing between an adult 
of this genus and a larva. 

In 1944 1 was able to make records of oviposition 
by adults of 0. swani n. sp., collected in north 
Queensland, and to rear larvae from these eggs, The 
present paper describes these experiments and also the 
adult, larva and ovum. 

Since the great majority of taxa among the Leeu- 
wenhoekiinae and other chigger mites of the family 
Trornbiculidae have been based on the larvae, one of 
the reared larvae is selected as the type of the species. 

I have taken the opportunity to examine the 
laxonomic status of some species of the genus from 
Queensland and New Guinea, and to give a key to the 
larvae of Odontacants known from New Guinea and 
South-Bast Asia. However it should be pointed out that 
this paper docs not undertake a general taxonormc 
review of the larvae of Odontacarus subgenus 
Leogontus, but considers only what is required to 
establish the new taxon proposed, and for the majority 
of the species considered in the key it relies on the data 
of previous authors. 

One species of adult allotted to this genus by 
Wumersley (1945) is markedly distinct from the others, 
ihis being O- retentus (Banks, 1916), and for it a new 
genus Scopitrombium is proposed, 

Revised keys are given for the classification, for 
adults and nymphs, of the subfamilies of the Trorn- 
biculidae, arid the genera of (he LeeuwenkockiirtMi 
All measurements are given in micrometres (^m) 
unless otherwise stated. 

Odontacarus Ewmg 


(Note: Only a partial synonymy is given here. For 
earlier and more complete lists of synonymies, see 
Wharton and Fuller (1952), Audy (1954), Goff and 
colleagues (1974-1979)). 

Odontacarus Lwing, 1929b, p. 188; 1931, p. 6; 1946, 
p. 436. Fuller, 1952, p. 228. Wharton and Fuller, 1952. 
p. 103 (ad part.)- Vercammen-Grandjean, 1968, p. 120. 
Loornis and Crossley, 1963, p. 381. Nadchatram, 1963. 
p. 535. Southcou, 1973, P- 46; 1976, p. 139; 1978, p. 16. 
Reed and Brennan, 1975, p. 6. Goff and Loornis. 1977. 
p. 370. Goff and Brennan, 1978, p. 50. Goff, J979a ; 
p. 143; 1979b, p, 140 

Leeuwenhoekia (Oudemans, 1911) Hirst, 1925, 
p. 150. Womersley, 1934, p. 217; 1944, p. 103. 
Womersley and Heaslip, 1943, p. 141. Taylor, 1946, p. 
227 ff. 

Acomatacarus Ewing, 1942, p. 490. Womersley, I94*S > 
p. 98. Taylor, 1946. p. 225. Grecnberg 1951. p. 525: 1952. 
p. 473. Fuller, 1952.. p. 229. Gunthcr, 1952, p. 39. 
Wharton and Fuller, 1952, p. 96. Audy, 1954, p, 164. 
Baker et ai 3 1956, p. 104. Dornrow. 1956, p. 150. 
Southcott, 1957, p. 146- 1973. p. 103; non Vereammen 
Grandjean, 1968. 

Odoniaairu* .vwar/i/ n. sp. 

Description of adult female (principally from 
specimen ACB240A, slide-mounted, somewhat com- 
pressed; but supplemented by other specimens) 
(Figs. IA-E; 2). 

Colour in life red. Idiosoma (Fig. IA) of normal 
ovoid shape for a trombidioid mite, without waist, and 
without division between prosoma and opisthosoma; 
length on slide about 1400 /un (ail measurements in 
micrometres), width about 900. 

Prosoma carries dorsally a well-chitinized crista, with 
an enlarged posterior end, carrying two sensiilary setae 
(Fig. IB). The crista tapers anteriorly, to a blunt- 
pointed rod, embedded in an arrow-shaped area of 
chitinization (here named the "sagitta* 1 since the term 
'tectum" used by Crossley (I960) for ihjs part of the 
scutum is considered less appropriate.). The crista is 
surrounded by the normal dorsal setation, sparser 
around anterior end. The sagitta carries two long. thin, 
filiate setae (Figs. IB, IE). Crista *43 long from its 
blunted anterior point to rear edge of the posterior 
sensiilary boss; 421 long to tip of sagitta. Behind 
posterior sensiilary boss the crista continues for a 
further 70, to end in a blunted point. Sensiilary boss 
94 across, distance between centres of sensiilary setae 
bases (i.e. **SB*) 57 Sensiilary setae slender, long. 


REC. S. AUST. MUS. 19(12); 169-200 

June, 1986 



FIG. L Odontacarus swani sp. nov. AduJi female, specimen ACB240, pareni of the hololype. A Entire specimen, slide-mounted, seen 
in transparency. B Crista and adjacent area. C Dorsal setae at posterior pole of idiosoma. D Lett palp, medial aspect. E Chclicerae, 
(All to nearest scale.) 

filiform, 290 long. The chitin of crista, including that 
of the area between the sensillary sockets, punctate. 
The whole of dorsum of idiosoma is densely covered 
with setae mounted on short papillae. The setae lend 
to be separable into two distinct types — a shorter burr- 
like, spindle-shaped, pointed, heavily ciliate group, 

mostly 27-31 long, and interspersed among these are 
longer, swordlike setae, in which the proximal part of 
the scobillum is somewhat expanded and ciliate; but 
the distal half is curved, swordlike, curved and pointed, 
with adnate cilia along the edges: a batch of these setae 
at the posterior pole of the idiosoma is illustrated in 



Fig. IC. There is, however, some intergrading between 
rliisv two rftafn typed of dorsal idiosomalae; ai the 
posterior pole the longer swordlike setae are 64-73 /mi 
long, the shorter spindle-shaped setae 25-42 long. At 
the centre of the dorsum the longer setae are 56-95 
long, the shorter setae 27-44 long. 

Eyes are not visible in any of the specimens of adults 
available, and it must be concluded that they are absent 
(despite being present and normal in the larvae). 

Ventral surface of idiosoma is covered with a dense 
selation of spindle-shaped, tapering, heavily ciliate 
setae. These tend to be uniform, similar to the shorter 
dorsal idiosomalae, except towards the posterior pole, 
where they are interspersed with swordlike setae similar 
to Ihose of the adjacent dorsum. 

Genital aperture in usual position, about 390 long 
by 2^5 across, with three oval suckers along each side. 

Anus normal, 91 long by 46 across. 

legs long, but robust for a trombidioid inr_c, 
moderately chilini/ed; trochanters generally rounded, 
b;t .ifemoral, iclofemoral and genual segments 
presenting as truncate cones, tibiae and tarsi approxi- 
mated cylindrical, leg lengths (including coxae and 
claws): I 2660, II 1760, III 1770, IV 2640. Each tarsus 
bears two claws. Legs well covered with numerous fine, 
pointed, lightly ciliate setae (scobalae), these being 
interspersed m Ihe distal segments with pointed, 
specialized sensory setae (spinalae, or eupathidalae). 

Tarsus I an elongate oval, 634 long by 214 across, 
wider than tibia I, which is 534 long by 178 across; 
tarsus II 397 long by 93 high; tibia II 313 long by l>l 
high; tarsus 111 399 long by S5 high; tibia 111 352x128 
similarly; tarsus IV 523 114 similarly; tibia IV 627x157 
similarly. For other leg morphometric data, see Table I. 







Crisla length' 









23 S9 





4fi 36 










32 4S 





72 91 


506 627 








53 27 

\ trout Jii!c-ti..ii cud 01 LlutMM'iiv rotJ M (Misterlor pok pI 

Omimns rfaws »n4 ptdtclc 

Palpi comparatively slender, well covered with ciliate 
setas (sec Pigi ID). Many elf these scobalae along the 
dorsal (i.e. extensor) margins of the palpal genu and 
tibia arc ciliate along their distal and outer aspects, the 
cilia broad and flattened, curved, uniform and almost 
apposed, as though capable of being touch receptors 
Utilizing n mechanism akin to stridulation; for these 
setae the term pectinate is proposed (plural; pectinalae). 
Palpal tibial claw stout, blunted, with four accessory 
claws (thickened scobaku ) on the medial to dorsal 

Palpal tarsus an elongate ovoid, with numerous 
ciliate setae (scobalae); these setae tend to resemble the 
palpal tibial and genual pectinalae, but are shorter 
Among them are also nude setae (spinalac) and pec- 
tinalae. A cluster of short sensory >etae is present at 
the tip of the palpal tarsus. 

Chelicerae robust; fangs curved, pointed, about 135 
long, with a row of about 10 fine retrorse teeth along 
distal flexor edge; extensor edge smooth. Chelicera 
basis about twice as long as fang. Rostrum (cone) of 
moulhparls with numerous ciliated, tapering setae, 

Taxonomic placing: In the key of Womerstey (1945, 
p. 110) this adult keys to the caption for O. tlron>>>: 
(Worn., 1939). From that species O. swatti may readily 
be separated as follows; 

Colour of adult mite red. Longer dorsal idiosortial 
setae with long cilia in the basal half, and with adttatc 
cilia, scarcely detectable, in the distal half of the seta. 
Shorter dorsal idiosoma! setae strongly ciliate, spindle- 
shaped, tapering . O. swani n. sp. 

Colour of adult mite white in life. Longer dorsal idto- 
sornal setae ciliate throughout length. Shorter dorsal 
idiosomal setae slender, uniformly ciliate 
O. dromus (Worn.) 

Further comments on the taxonomy of adult iceu- 
wenhoekiine miles are made later in Uiis article, 

Description of Ovum 
(Fig. 2, 3A-D) (from mounted material) 

Colour in life not observed. Ovum nearly spherical 
to somewhat spheroidal, about 250-350 long in longest 
diameter. Ovum has the usual lightly pigmented outer 
layer (chorion), which is smooth, lacking any striate 
markings in the available material. Minute tuber- 
dilations occur on The parts of the mounted material, 
but these could be, 3t least to some extent, result* oF 
compression in eciosion, or rupture by other means. 

I he sole embryo available (Fig 3D) is about 280 long 
in its greatest diameter, by about 220 wide in its shortest 
diameter. It contains a large number of rounded 
granules, presumably of yolk, mainly concentrated in 
the region opposite to that in which the limbs are 
developing. TWo annuli (seta bases) are present in the 
specimen, and two tooth-like structures, possibly shell- 
piercers in eciosion ("t" Fig. 3D). From one annulus 
there is a vague indication of a developing seta shaft. 
On the developing limb segments are smooth, rounded, 
low tubercles, similar to those recorded for Trornbcih 
alpha developing nymph (Southeou 1986a, Fig. 5A>. 

One chorion (Fig. 3C) is unusual in being covered 
randomly with small circular or oval lacunae In 
rupturing, presumably from eciosion, some of these 
lacunae have been cut across transversely; it is clear that 
they were present before rupture, and presumably in 
life. No evidence of mould filaments or other possible 
chilmolytic agents can be seen near them, and their 
origin is unknown. Under polarized light a few of them 


REC. S. AUST. MUS. 19(12): 169-200 

June, 1986 

FIG. 2. Odontacarus swani sp. nov. Adult female, specimen ACB245, seen in transparency. The idiosoma contains nine eggs, of varying 
sizes, some having been disrupted by the mounting process. 



FIG. 3. Odontacarus swani sp. nov. Ova, slide-mounted specimens. A Ruptured ovum, showing chorion. B Chorion of another ovum 
split into three pieces, and possibly some of the developing embryo. C A ruptured chorion with multiple lacunae (sec text). D Developing 
embryo, shed from its chorion, with early development of limbs and other structures: a annulus or seta base, t tooth-like projection 
(see text). All specimens shown in transparency. (All to scale shown.) 

show a typical Maltese-cross figure, indicating that 
there is possibly a layer of optically active material at 
the base of the lacuna, perhaps an artefact of the 
mounting. The chorion shows only weak evidence of 
optical activity under crossed polarizers. 

Description of Larva (from reared specimen 
ACB240B, holotype, and supplemented by other spe- 
cimens) (Figs. 5A-D, 6A-C). 

Colour in life red. Length of idiosoma (mounted on 
slide) 207, width 197; total length of animal from tip 
of cheliceral fangs to posterior pole of idiosoma 293. 

Dorsal scutum slightly wider than long (including 
nasus). Although most authors measure the length of 
the dorsal scutum ("S.D." or shield depth) of 
Odontacarus larvae (and other genera of the Leeuwen- 
hoekiinae in which there is a distinct beak-like "nasus" 
to the scutum) excluding this "nasus", a comparative 
study of allied forms, e.g. Neotrombidium larvae, as 
well as larval Trombella (see Southcott, 1986a), shows 
that the "nasus" may range from a broad continuation 
of the scutum, without clear point of differentiation, 
to a distinct and sharply demarcated beak. The length 


Kf I . S. MIST. MUS. 19(12): I69-2(K) 

June, 1986 


FIG. 4, Conventions of measurements of dorsal scuuim of B tecu- 
wenhockiinae larva, as used in the present article (sec text). 

(L) is taken here as including the u nasus". However, 
in order to make the measurements comparable 
between different workers, measurements are offered 
here of the shield length including u nasus" (L) and 
without "nasus** (LB) (see Fig. 4 for explanation), 

Nasus well-developed, running into more or less 
straight (slightly sinuous) anterior margin; lateral 
borders slightly convex; posterolateral convex, of 
varying curvature; AM setae tapering, pointed, lightly 
ciliate; AL and PL similar. Scnsillary setae normal, 
ciliate in distal half or third. Sensillary sockets about 
level with PI. bases, sloping obliquely anterolateral^, 
and with well-marked margins ("lids"). 

The standard (and other) data for Ihis specimen, as 
well as other reared material, and the Type series and 
olher identified material in the South Australian 
Museum Colled ion are given in Tabic 2. 

Eyes 2+2, sessile, near PL angles of dorsal scutum, 
oval. Anterior eye with maximum diameter of cornea 
16, posterior 14. 

Dorsal idiosomal setae long, moderately ciliate with 
lightly projecting barbs; setae arranged 6, 12, ..., in 
vague transverse rows, lotai |fl AC B240B, 57. 

Venter with a pair of pointed setae between co\ac 
III*, ciliate, 33 long. On opisthosoma ventrally about 
37 further setae, pointed, well ciliate, 27-42 long, the 
posterior setae longer and lending to be blunted at the 
tip, resembling the posterior dorsal idiosomalac 

Coxalae 2, 1, I- All coxalae long, pointed* ciliate; 
lateral coxala 1 51 long, medial 46, coxala II 44, (II — 
(missing in holotype; 58 long in ACB240C ). 

Legs normal, I 379 long, II 322, 111 380 (all lengths 
include coxae and claws). Chaetotaxy of legs as figured, 
Leg scobaiae normal, pointed, ciliate. Trochanteral 
Kumula I, I, 1; femoral 6, 5, 4. Leg specialized setae 
as follows; VsGeL70pd ("microseta"), VsTil.87pd 
C'mieraseu"). SoGcL.lftpd, SoTil.90d (i.e. slightly distal 
to Vs), VsGelL70d, SoGcl!.30d, SoTill.45d, SriTiIL9ld, 
SoGellt.29pd. Tarsus 1 and II have each a large dorsal 

1 hutacter 












P ft 
































17 32 









4 4^2 

I N 








B5 ''7 




A -I 1 


























II a 


12. 00 







PW ! I* 1 

1 22 






( 3<v44H 47-51) 





56 44 



46 25- 







1 727 J 











55 .25 



















1 581 







this is the PW/SD of Womcrslcy (1944. 1945] ttrtd ihf PW/L of 

Greenberg (1951) (sec- I ig. 4, and text explanation!. 
: For the maximum values of each variatc. 

solcnoidala. FaTall present (see figure). Normal SoTalll 
present. Tibia 111 with two mastisetae (=masialac): 
MaTilll.73d, MaTiilL76pd. Tarsus III with ft 
rnnMiscta: MaTitll.33pd. (In these estimates tarsal 
length excludes claws and pedicle.) tarsus 1 86 long by 
32 across. Tarsus III 85 *24. Other specialized leg setae 
as figured. Other leg measurements as in lable 2, 

'ftirsal claws normal, falciform, slender, with strong 
cilia (onychotrichs) on anterior and posterior; middle 
claw longer and more slender than the neolaterals, with 
weak cilia. 

Gnathosoma normal. Combined chelieerae bases 84 
across, chclicerae 89 long from tips of fangs to 
posterior pole of chelicerae bases. Cheliceral fang stout, 
CUTVGCl, pointed, with three of four retrorse teelh along 
(concave) flexor (dorsal) edge, and 4-5 tubercutalions 
or blunted teeth on (convex) extensor (vetiiral) edge. 
Galcala nude, 22 long. Gnathobasal setae (palpal 
coxalae) curved, pointed, well ciliate, about 24 long. 

Palpi compact. Palpal formula I, 1, 3, 8. with palpal 
formula B(b), N<?b), BNN. So + 7 (B or N), as figured. 
Palpal libial claw txifurcatc. No supracoxala to legs or 

Origin of Adult Material Studied 

[ collected ten adults in damp soil in rainforest at 
Ml Hypipamee, Athcrton Tableland, Queensland on 
26, x, 1 944. 

Each was placed in a small tube with some damp 
soil from the capture area, and observed periodically 
(albeit somewhat irregularly, as I was engaged in 
military duties). Living larvae were observed in two of 
the tubes from 20-24.xu 1944 using a 23* Zeiss 
microscope ocular reversed as a hand lens; in a third 



FIG. 5, Odontacarus swani sp. nov., larva, holoiype, ACB240B. A Dorsal aspect, legs on right hand side omitted beyond trochanters. 
B Cheliceral fangs and galeala from above (distal haJf of cheliceral fangs shown in transparency). C Posterior dorsal seta (x in A). 
D Posterior ventral seta (y in Fig. 6A). (All to nearest scale.) In this and in subsequent figures the A sign indicates the seta is shown 
in bolh dorsal and ventral figures. 


REC. S. AUST. MUS. 19(12): 169-200 

June, 1986 

FIG. 6. Odontacarus swani sp. nov., larva, holotype, ACB240B. A Ventral view, legs on right hand side omitted beyond trochanters. B 
Right palp, dorsal aspect. C Right palp, ventral aspect (B, C to scale on right). D Claws of leg II, further enlarged (not to .scale). 




Serial No. 





Adult dead 


A Cfi 










(S.vii 45)* 

20-23. xii.44 



























Gravid, laid no eggs, but 
body contains about 50 ejuii 






22.xi.44-20.Jui 44 




.— i 

-.a 6_m.44 

Oravid (9 tggs), laid noix- 





12 si.44-20.xii 44 



22.xi .44-20. xii .44 

• rhcse egg* or larvae were found iii the dned tubes, months or year? after the attempted rearmgs had been concluded 

tube examined some years later (long after all the tubes 
had been allowed to dry out), larvae were also found, 
dead and dry. Although living eggs were not observed 
in these tubes, egg remnants were eventually Found 
when the oppon unity to examine the tubes in detail 
with a microscope occurred. Two female adults, on 
being slide-mounted, were also found to contain eggs, 
although no eggs or larvae were recovered from their 
tubes. The details of these experimental animals are 
summarized in Table 3. 

Taxonomic Placing of Odontacarus swani n. sp. 

The larva of this species is distinguished from others 
of the genus by the following combination of 
characters; (1) palpal tibial claw three-pronged, (2) 
galeala nude, and (3) about 57 dorsal body setae 
present, (4) two mastitibialac 111 present, (5) one 
mastitarsala III present, palpal genuala (PaScGe) and 
the two palpal tibialae (PaScTil and PaSeTi3) almost 
nude, with only faint indications of barb.s, cheliceral 
fangs with a row of 3-4 dorsal and 4-5 ventral denticles. 

Characters (1), (2) and (3) alone are sufficient to 
distinguish this species from all others of the Australa- 
sian to South-East Asian region (subgenus Leogonius). 

The diagnostic characters of the adult have been 
given above. 

This species is dedicated to Duncan Campbell Swan 
(28.xi.l907-26.xii. 1960), entomologist, who made field 
observations on the distribution of chiggcr mites in 
South Australia (sec Southcott, 1982). 

Classification oj the Larvae of the Australian, New 
Guinean ami South-East Asian Species of Odontacarus 

About 20 species of Odontacarus have been 
described from Australia and New Guinea (including 
West Irian and Papua New Guinea). A single further 
species, O. audyi (Radford, 1946) has been described 
frurn South-East Asia, one, O. gymnodactyii (Ewing, 
1925) (1925b) from India, and one, O. tygosomae 
(Dumbleton, 1947), from New Zealand. Since only 
limited keys to the classification of these species have 

been offered since Womcrsley's (1944, 1945) papers, an 
attempt to key the species of this region is offered here. 

The most detailed studies of recent years for this 
region have been done by Goff and Loomis (sec refer- 
ences), who unfortunately deal only with Ihe species 
from New Guinea, and incidentally with a few north 
Queensland species, where these occur also in New 

In attempting to classify the larvae here allotted to 

0. swani n. sp M as well as various other species from 
northern Queensland and New Guinea, 1 have experi- 
enced difficulty in using one of the main character* 
used by Goff (1979a, b) in classification, this being 
deciding on the number of setae on the dorsum of the 
abdomen in the "first post-humeral row", i.e. the row 
of setae immediately behind the dorsal scutum, but 
excluding the two (or four) long "humeral" or 
"scapular" setae. These remarks apply not only to the 
specimens of O. swani described here, but also 
members of the type series of O. afhenoriensts 
(Womersley, 1945), and of Q mccutlochi (Worn., 1944), 
which were used by Goff (1979a) in drawing up his ke\ 
to the species which occur in New Guinea. 

For this reason, it has been decided not to use the 
number of the setae in this row as a key character. If 
should also be noted that students of the larvae of the 
Trombidioidea and Erythraeoidea usually experience 
difficulty in deciding which setae to allot to different 
rows, as the arrangement is commonly irregular, and 
the allotment of the setae is largely a matter of 
guesswork. On the other hand, the total number of 
dorsal or ventral idtosomal setae appears to be free 
from these defects, even if the counting of them i.s at 
limes tedious. 

The following key to the larvae of Odontacarus for 
this region is therefore proposed, based to a large extent 
upon already published descriptions, but supplemented 
with some new data. 




1. Palpal tibial claw with more than three prongs „ S 
Palpal tibial claws with three prongs -41 


Rf.C. S. AUST MUS. 19(13$ 169-200 

June. 1986 

2. Palpal tibial daw with seven prongs 

O riannaeGod, 1979 

Palpal tibial claw with four prongs 9 

3. AL setae longer lhan VI 

(I nudvhamnni Uifl, WW 

AL setae shortci than l*L 4 

4. Miisutihialae III absent Two rnastilarsalae 111 present. 

Dorsal idiosomal setae about 110 

O. mitchelli Gofl , 1981 

One or more rnastitibialae HI present 5 

5. One mastitibiala 111 present. Dorsal idiosomal Sfiiae 

about 230 O. unisetosus Goff. 1979 

Two rnastitibialae III present to tibial 111 ,.,f 

6. Galeala nude. Palpal selal formula 1)/B/DNN/So * 7B 
About 120 dorsal idiosomal setae present . 

. - O. Iriselosus Gofl. I^7V 

Galeala barbed .7 

7. Dorsal idiosomal setae number about 54 Ventral 
idiosomal setae about 78. Palpal setal founula B/B/ 

BBB/So f 7B. Cheliceral denticles 4-5d/4-7v 

O. zteglen Got!'. 1979 

Dorsal idiosomal setae uurabcr about 5,1-90. Ventral 
idiosomal setae number about 46-56. PaipaJ setal 
formula B/B/Bbb/ScM 7B. Ctieliceral denudes 7d/ 

6-8v O. audyi (Radford, 1946) 

(Occasional specimens of O audyi have only three 
palpal tibial prongs (see Nadehatram, 1963).") 

8. Mastitibialae III lacking 9 

MastitibiaUe III present ... J I 

9. Mastilarsalae 111 lacking. About 42 dorsal idiosomal 
setae present. Cheliceral denticles 3-4d/3-4v. AM seta 
short and thickened, reaching about 2/3 across to the 

AJL seta base (Fig. 7 A, B) 

O. soufheotti (Womersley, 1944) 

Mastitarsalae III present 10 

10. CfattOend denticles 4-5d/ 1 v (Fig. 7C). DS to 22 u,m long 

O. langarti (Southcott. 11*57) 

Cheliceral denticles 3-4d/4-5d (Fig. 7D). DS 29-32 ^rn 
long O. mathewi (SouthcetL 1957) 

11. Tibia 111 with three rnastitibialae. Palpal setal formula 
B/B/RBB'So<7B. Cheliceral denticles 3d/3v. One 
masmarsala m present 

O. mvcullovht (Womersley, 1944) 

Tibia IH with two rnastitibialae ............ I?. 

12. About 30 opisthosomal ventral setae present 

O. adelatdeae { Womersley, 1944) 

More than 30 opisthosom*! ventral setae present 13 

13- PL/AL less than 1 .20 

. O. nova-guinea Womersley, 1944) and 

O. fonxipes (Womersley, 1945) 
PL/AL greater than t.20 14 

14. Over 75 dorsal idiosomal setae present 

. O. ausfratiensi* (Hirst, 1925) 

and O hirsli ^ (Womersley, 1944) 
Less than 70 dorsal idiosomal setae present 15 

15. More than 58 ventral optsthosomal setae present. 

Cheliceral denticles 3d/4v (Fig. 7F.) 

.0. cooki (Sourheou T 1957) 

less than 58 ventral opisthosomal setae present . 16 

16. PSB/SB <1.00 1 . PW 87 ,mwl ,ou (Womcrslev, 1**45> 

. i. . O. tMJrhnensis (Womersley, 1945) 

PSB/SB al.OU 1 . PW mostly less than 85 /mi ... , , 17 

17. PL/AL > I. -V Ventral opisihosomal setae 48-54 In 
number Palpal setal Formula B/B/BBB/So + 7B. .. 
....... ... O. athertonenm (Womersley, 1945) 

PL/AL < 1.3. Vcnrral optsthosomal setae En range of 

35-45 in number - - 18 

1 For O. burrinensis and O. eehutttus this statistic is 
based on Womersley's (1945) Figs. SF and 5D respec- 

18. AW 54-6). PW/(A-P)<2 75 : 

O. echidnas (Womersley. 1945) 

AW 65-73. PW/(A-P)>2 75. Palpal setal formula 
B(b)/M?b)/BNN So i 7|B or N F . Galeala nude . . 
O, Swunt n. Sp. 

The following two species have been omitted from 
the above key: 

(1) Odontacarus gymnodanyh (Ewing, 1925) 
(1925b), described from India. This species was 
doubtfully referred to Acomatacarus (Acomatacarus) 
by Wharton and Fuller (1952, p. 98), who list the 
previous synonymy. The descriptive details aie 
insufficient to place this species in the above key. 

(2) Odontacarus tygosomae (Dumbleton, 1947), 
from New Zealand. The descriptive details are 
insufficient for the placing of this species in the above 
key. Nevertheless it is quite a distinct species, with a 
wide dorsal scutum, and flattened, broad, ianceoiate 
scutalac and dorsal idiosomal setae, the latter 
numbering about 100. 

Analysis of Morphometry Data of Q. swani Larvae 

In previous papers ] have attempted to determine 
correlations and other relationships between variates 
that are used by taxonoraists for dassificatory 
purposes, to see if they may be a useful basis for t0QJ£ 
general studies such as whether the mechanisms 
cvmtrolliri£ si*e are interrelated in some way, e.g. 
genetic Southcott (1966, pp. 736-738) reported on the 
correlation of the standard data for the erythraeoid 
larval mite, Charletonia swaiiana (Lawrence, 1940). In 
a later paper (Southcott, 1 986a) a coi itrlation study by 
principal component analysis (Hotetling, 1933) was 
presented for both seutal and leg variates for a 
trombidioid larval mite, Trombella alpha Southcott. In 
the present paper a series of larval O, swani is similarly 
analysed. Some of the same remarks made about 
utilizing the data or Southcott (iy86a) apply also to the 
present paper. In attempting to elucidate relationships 
we have here a group of larvae reared from conspecific 
adults, collected over a few square metres of soil, near 
the surface, on a single day. Additionally, the batches 
of larvae were reared with damp soil jn sealed tubes 
in my baggage, so that all the larvae of each batch may 
be said to have experienced at least similar conditions 
in their life-spans. Such conditions could therefore 
possibly eliminate sources of variation from dissimilar 
conditions of rearing. 

In making statistical analyses, as was done with (he 
study of the larvae of Trombella alpha, we may ftQriCI 
the data for the dorsal setae to a single variate, in view 
of the partial redundancy in using the variates DS t mid- 
DS and PDS. We may also ornit the data for Sens, since 
these scnsillary setae are slender, flexible and difficult 
to measure accurately; also these setae are commonly 
absent in microscopic mourns, whether from the 

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RbC. N\ AtJST. MUS. 19(12): 169-200 

June, 1986 

capture, mounting or preservation procedures 
themselves or possibly earlier traumas of life. Omitting 
also PW/LB, which is a statistic derived from two of 
the listed variates, we may therefore construct a 8 > 23 
table of variates from these data. 

A correlation matrix of 23>;23 values (including L, 
which is the sum of LA ¥ LB, or ASB + PSB) was then 
calculated. There are only 21 independent measures in 
the table of r, on 6 degrees of freedom (Table 4». 

The table of the correlation matrix shows a total or 
25 significant correlations (at P=.05 or at greater 

From this we may calculate a principal component 
analysis (Table 5). As there were only eight specimens 
in the series, it was considered the principal component 
analysis should be restricted to only two (first and 
second) components. 

Table 5 for the principal components for the variates 
selected for study in O. swani shows that the first 
principal components includes all 23 vacates with the 
exception of SB, PSB, LA, LB, A-P, AM, PDS, Til, 
Cell and Till, indicating that this component may 
represeni a size variable affecting the majority of the 
variates; and the second principal component includes 
PSB, L, LB, and AM (also LA and Till, of opposite 
sign of the direction cosines), thus indicating that these 
measures are correlated, and that after allowing fot size, 
where one group is relatively large, the other is relatively 

Further Studies on the Relationships of Variables 
Used in Classification of TYombicalid Mite Larvae 

Statistical analyses of chitinized parts of various 
Irombidioid, erythraeoid and other mites have been 
increasingly used as an aid to classification. My use 
of correlation analyses to determine the interrelation- 
ships of these variates has been outlined above. In my 
J 986 (1986a) study I showed that there is a correlation 
between a variate dependent upon a body character 
(scutum) and of a leg character, in some cases. Above 
it was found that in O. swani larvae both principal 
components (first and second) include both scutal and 
teg dimension variates, these being positively correlated 
in the case of the first, and both positively and 
negatively (for the different variates) in the case of the 

In OTder to decide whether such interrelationships 
are an expression of some more general phenomenon 
or phenomena* it was decided to do similar studies on 
some other species of related trombieulid mites. 
According to keys in existence befoie the present study 
was commenced, it was thought that the larvae 
described above as Odontacarus swani were classified 
close to Q athertonensis and O. mceullochi, and 
initially were classified as O. athertonensis. Both of 
these species are included in the same couplet in the 
key of Goff (1979a, p. 154). These two species of larvae 
were therefore selected as being suitable for similar 

correlation studies, with the additional hope that if 
difficulties in classification were to appear, the 
correlation study might provide some guidance as to 
the relationships between these species. Accordingly 
careful measurements were made on the available series 
of these two species also. Ultimately, with further study, 
it was found that the larvae described here as 0, swunt 
differed in several characters from both O. 
athertonensis and O. mcculbchu so that only O. 
athertonensis and O. swam appear close together in the 
key above. 

In the case of O. athertonensis a long series of larvae 
was available (28 specimens), and in the case of O. 
mccutlochi, 10 specimens. These were measured for Lhe 
same structures as in Q swani Nevertheless, it is clearly 
as valuable to test for species widely separated from 
each other as for closely related species, in a search for 
such interrelationships. 

OdotiUcvrus athertonensis (Womersley) 

Aeomatacarus athertonensis Womersley, 1945, p. 104; 
Taylor t 1946, p. 226; Fuller, 1952, p. 230; Gunther, 
1952. p. 40; Wharton and Fuller, 1952, p. 97; Audy, 
1954, p. 164, 

Odontacarus athertonensis Goff, 1979a, p. 144. 

Source of the Larvae of Odontacarus athertonensis 
Used in the Present Study 

The larvae of Q athertonensis studied for the present 
paper came from the following sources: 

(1) The type series of 19 larvae, identification 
numbers ACB732A-S, Wongabel, Q., October, 1944, 
R. N. McC[ulloch], also two further larvae, identi- 
fication numbers ACB732T, U. with same locality, date 
and collector information; South Australian Museum 
collection. Specimen ACB732A slide is notated 
"Syntype"; specimens ACB732B-S arc notated as 

























- 0.2846 
~ 0.2688 
0.31 1 1 
0.1 108 
■ 2384 
i) 2891 
o is?: 


II 0850 



O 1144 
o 3843 



Paratypcs; specimens ACB732T, U are without type 

Womersley stated (1944, p. 105) that the species was 
'•described from 16 syntypcs n . Specimen ACB732A, 
which bears also a number N1981384, and was 
presumably the one examined by Goff (1979a), is 
hereby formally designated the lectotype* 

(2) Six larvae, mounted individually on slides, 
identification numbers ACB733A.-F, Mt. Jukes, 
Queensland] [2I°00'S. 148°57 El, 6.ix.l95I, E. R. 
Derrick (South Australian Museum collection). These 
six slides were also examined by Goff (1979a), and bear 
the notations 0192, GI93, G197, G196, G206, G197 
respectively, also numbers GI981385-GJ98I390, 
respectively. Each slide also bears the comment "Was 
marked paralype" (this marking has been removed). 
These slides were clearly never part of the type series. 
Specimen ACB733C is a poor mount, unsuitable for 
detailed measurement. 

Omitting specimen ACB733C, three erf the five 
larvae key directly to O. athertonensis by the key given 
above, while lor two others (ACB733 E & F) O. 
athertonensis is the nearest fit, (here being a mismatch 
in each of PI./AL- 1.225. Further studies of these 
larvae have been presented elsewhere (Vcitch and 
Suuilieott, 19K4; SaulhcuU, 1986b). 

(V) ACB734, one larva, Lae lest Area, Papua New 
Guinea, 24.iv.l944, R. N. McCulloch (S.A. Mus.). The 
slide has the notation; Leeuwenhoekia (Acorrtatacanjs) 
nr. athertonensis Worn. Lae Test Area. 24.4.44. R. N. 

McC. Det. H. Womersley. AW and PW Standard Data 
longer. AW = 75.6, PW-84.0. (This is a poor mount, 
the dorsal scutum being damaged.) 

By the key given above this specimen comes to 
caption 18, but then does not fit the criteria of either 
O. echidnas or O. swani. A further analysis of (he 
statistical data of (his specimen has been made 
elsewhere (Vcitch and Southcotl, 1984; Southcott, 

In response to my request for locality data, Dr. H- N 
McCulloch has replied (pers. cornm., 1983): "Lae test 
area was jungle just off'the main road and where 1 did 
stopping-time tests [see McCulloch 1944, 1946, 1947] 
probably a mile towards the big air strip from 2/7 
A, G. H/' (2nd/7tb Australian General Hospital). 

(4) ACB735, one larva, Lae Scrub, 24.iii.44, A. S. D. 
(South Aust. Mus.). The slide is additionally notated: 
''Leeuwenhoekia (Acomatacarus) nr afhertoncr.-, 
Worn. AW and PW Standard Data longer. AW=70.0 f 
PW=84.0. Det. H. Womersley." This is another poor 

By the key above this specimen Is not identifiable, 
owing to both legs III being damaged 

As no mite student or collector with the initials 
A. S. D. was known to me, I have written to Dr. R. N. 
McCulloch asking for further locality and other 
information. He has replied (pers. comrn., 1983) "ASD 
means, 1 guess, Advanced Stores Depot and was on the 
old disused civilian air strip which was between the new 
[i.e. 1944] active air strip and 2/7 A. G. H." 



/ ec to type 




Range for 
Type series 







61 71 



















1 413 




54) 52) 4 








2806) 4 








K2(K8) 4 



M 90 









I El 



M(70> d 












































2. 101 























nw l B 
















mid DS 







2 385 1 



42 4fi 

tt-42(-S)) i 




1 964' 








1 .670 







< e iy 


Tic 11 


















42(44) J 




45 71 










1 The mean and the s.d. arc calculated for tht type series only with n 21 (see text). 

: In subsequent calculations ot correlation the three missing values indicated by (a), (b) and (c) were replaced as follows: (a) At lor 

ACB732B: 41, (hi AM lor ACB732S: 35, <c) Til for ACB7J2B; 53, after a study ot the data. This refers to the table of vanates Tor 

the type series held on lile in the South Australian Museum. 
1 figures calculated from the maximal values only ot that vanate. . 

* figures in brackets are S set of variant estimates made subsequently and which differed by 2 ,itn or more trom the earlier ones, and 

used |n JalculatiofW prcattited in Veiteh and Southcott U9K4) as a check on the reliability of analytical procedures. I he specimens were rlfC 

m08l rtparflUJcl from the others using the variables selected. Specimens ACB734 and ACB735 svere poor mounts, presenting difficulties jn 

microscopic measurement 


REC. S. AUST. MUS. 19(12): 169-200 

June, 1986 



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A further statistical study of the data of this 
specimen has been made elsewhere (Veitch and 
SoUttKOtti I9S4; Southcott, 1986b). 

(5) ACB736, one larva, collection data as for 
ACB734. The slide bears the notation; "teeuwenhoekia 
(Acomatacarus) nr. athertonensis Worn. Lae Test Area. 
24.4.44. R. N. McC AW and PW Standard Data 
longer. AW=68.6, PW=84.0. Det. H. Womersley." 

Remeasuring gives AW about 65, PW about 75. 
From obscurity in the mount the specimen is not 
identifiable by the foregoing key, various key characters 
not being visible. (The data of this specimen have been 
omitted from the table given below (Table 4) t as too 
many values are unavailable for it to be capable of 
being included in a correlation analysis.) 

These three are the only specimens in the South 
Australian Museum collection placed as or near O. 
athertonensis, from Papua New Guinea. Although 
Goff (1979a, p, 146) states that the three specimens 
examined were from "Morobe Distr[.]> Lae, 24.iv.1944 t 
collected in scrub", only two slides bear the date 
24.iv.1944, and the term "scrub" occurs on only one 
slide; none of these slides carry any loan number 
designations, apart from that implied by my 
identification numbers. 


The measurements of the standard data and other 
variates selected for study are shown in Table 6. 

Results of the Correlation Analysis of the Variates 
o/Odontaearus athertonensis Larvae 

)n Table 6 the means and standard deviations of the 
variates have been calculated from the type series only 
(ACB732A-U). The reason for this is that a preliminary 
discriminant analysis of the data given indicate that 
the specimens previously considered as conspecific, as 
well as those under the heading "Conspecificity 
queried" differ significantly in various statistical 
measures from the type series. This subject will be 
analysed further in another paper? Accordingly, 
correlation analyses here will be confined to the data 
of the type series. 

Proceeding as before, a correlation matrix of 23x23 
values is calculated; this is shown in Table 7. 

There are 32 correlations between variates shown in 
Table 7 at the 5 a /o (or more significant) levels of 
probability. These are marked in the table by asterisks, 
as shown. 

Examining the principal components as before, we 
may select the first and second principal components 
(in view of the sample size of 21 specimens). These are 
shown In Table 8. 

From Table 8 wc sec that there are two significant 
groups of correlations. Component I, of AW, PSB, LB, 
Til, Gel J, Till, Gelll, Tilll, together with (of opposite 

* Note added in proof: This has now been done; see Veiich and 
Southeott (1984). 










I B 











i i 




• 0.3569 


sign) ASB, LA, LN and A-P. This component thus 
represents a mixture of size and shape variables. 
Component 2, of ASB, PSB, L, LA, LB, LN and PDS, 
and (of opposite sign) A-P, which is thus seen to include 
mainly variables based upon the dorsal scutum. This 
component again is a mixture of size and shape 

Odontacarus mcculiochi (Womersley) 

Leeuwenhoekia mcculiochi Womersley, 1944, p. 108. 
Acomatacarus mcculiochi Womersley, 1945, p. 111. 

Tkylor, 1946, p. 226. Gunther, 1952, p. 40. 
Acomatacarus (Acomatacarus) mcculiochi Wharton 

and Fuller, 1952, p. 99. Audy, 1954, p. 164 
Odontacatus (Leogonius) mcculiochi Goff, 1979a, 

pp. 147, 154. 

Material Examined 

(All specimens in South Australian Museum 


Trinity Beach Area [Trinity Bay] 10.vii.1943, R. N. 
McQuIloch], on boots, edge of scrub, four specimens, 
identification numbers ACB752A-D, A labelled Type, 
B-D Paratypes. 

Trinity Beach, Cairns, Qfld.], edge of scrub, 
8.vii.l943, name of collector not stated, identification 
numbers ACB753A-D. (These slides are labelled 
Paratypes, although according to Womersley *s account 
(1944, p. 109) they were not mentioned in the original 

Emerald, Qentral] Queensland), Nov. 1948, ex 
Pomatostomus temporalis, H . W (omersley L two 
specimens, identification numbers ACB754A, B. 


The measurements of the standard data and o>hcr 
variates selected for study are shown in Table 9. 


K£< s. AUSTt MUS. 13(12)! 169-200 

June, 1986 








S 17. 




































A 358 

















26- 30 

4 T 13fl 

















35 <>0 

2 169 












3 65l 





10 90 

! 28T 







J. 38 















34.70 1 






■*8 70' 

2 406' 















J* tf 


42 20 

2 741 









4^ 54 


4K 00 


Til II 



1 ' 



I. Figures for the maximum valuer o! tfcfc vftl 

Analysis of Morphometric Data for Odoiuae.;.in's 
mccullochi larvae 

The data in Table 9 were submitted lo the previously 
described analysis, the variate Sens being omitted tor 
the reasons as before. Likewise the variates involving 
the dorsal idiosomal setae were restrieied lo the 
maximum values of the mid-DS and PDS variates. 

Proceeding as before we thus have a 23 x 23 table on 
8 degrees of freedom (T&ble 10). 

Table 10 shows 28 significant correlations between 
the variables, at the 5% or greater level of significance. 

From these data wc mav perform a principal compo- 
nent analysis, as before. This is shown in Table II. 

From Thble II il can be seen that there are two 
significant groups of correlations. Component I, of 
AW, PW, SB, ASB, L, LA, BL, LN, W, A-P, AMB, 
Gel, Til and Till, i.e. indicating largely that this is a 
eoi relation induced by size factors. Component 2, of 
AW, Til, TiJl, Gel II and Tim, and (of opposite Si 
LA, LN, A-P, AL, AMB, MDS and PDS; it isthu 
mixture of correlations of opposite signs, involving 
variables derived from scutum, body setae and leg 

Conclusions on Correlations He? ween Measurements 

ufChittnizect Idiosomal and Leg Parts in Three Specie* 

of larval Trombidioid Mites 

My earlier study on Ttomhella alpha {SuuiheutL 
1986a) and ihc picsent study ol Ihree species Of 
trombiculids have defined in each species the first two 
(or three) principal components in a comparable series 
ol vai iates from specified chitinized parts of the body 
and limbs ol these mites. The findings are set out in 
Table 12. 

It is apparent from the preceding analysis and Table 
12 that the correlation patterns are different in each 
species of mite studied, for the variates that have been 
selected for siudy. For the first principal component, 
there appears to be a greater similarity between the 
patterns between O. mccullochi and O, swani, than 
between either of these species with O. athertonensis. 
The patterns of the first principal component in these 
three species of Odonracarus resemble each other more 
than they do that of Trombella alpha, 

Tins result suggests that all of these measurements 
may be necessary to separate species of Odonfucants 
and, by an extension of this concept, species in other 
genera of mites. 

There is thus not a compact sei of correlations mani- 
fested in these data. This is due in part to the small 
sizes of the samples studied, although other causes of 
these variations and comparative paucity of correla- 
tions could be suggested, particularly nutritional or 
other environmental factors. 

Odotitacarus saulhcotU (Womersley) 
(Tig. 7A. B) 

Leeuwcnhockia southcotti Womersley, 1944, p. 109. 

Tayloi, 1946, p. 232. 
Acomatacarus southcotti Womersley, 1943, p. 111. 

Taylor, 1946. p. 226. C.unlher, 1952, p. 40, 
Acomatacarus (Acomatacarus) southcotti Wharton 

and Fuller, 1952, p. 100; Audy, 1954, p. 164. 

Material Examined 
ACB169A1-7, South Australian Museum collection, 
Adelaide River. I5.V1.1943, on skink. (These are the only 
specimens in the S.A.M. collection; originally J had 
submitted 10 specimens, on two slides, labelled 
ACB169A (8 specimens) and ACB169B (2 specimens). 

Remarks on Collection Data 
On I collected 10 or more red larval mites 
from the external auditoiy meati of a small skink, 
specimen R45 (my number), in the Adelaide River area, 
about 12 km (8 miles) east of the 57 mile point <?jk. 
57 miles (91 km) southwards along the main road from 
Darwin; this being the current military nomenclature). 
This is about lat, 13 06 S, long. 131°14'£ ( There were 
5 or more larval mites in each ear of the skink; but 
none in the axillae or elsewhere on the external surface 
of the skink. The skink was later identified as 
Uiolcpisma ?pectoralis by Mr F. J. Mitchell, South 
Australian Museum, but not kept. Another specimen 
caught at the same lime and place (my R46) was 
identified by Mr Mitchell as L. pectoratis, and is 
preserved in the South Australian Museum colled ion 
as R2703. 

These notes supersede the collection data given by 
Womersley (1944, p. 110), which are partly wrong, 












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June, 1986 




























- 0.0299 

Additional Notes on Morphology 
This species has a broad, shallow dorsal scutum, with 
short, rather thickened, AM setae, which reach about 
two-thirds of the way across to the AL seta bases. The 
eheliceral fang has 3-4 dorsal, and 3-4 ventral, denticles. 
These features are illustrated in Figs. 7 A, B. 

Odontacarus barrinensls (Womersley) 

Acomatacarus barrinensis Womersley, 1945, p. 106. 

laylor, 1946, p. 226. Gunther, 1952, pp. 41, 48. 
Acomatacarus (Acomatacarus) barrinensis Wharton 

and Fuller, 1952, p. 98. Audy, 1954, p. 164 (with 


Remarks on Collection Data 
Womersley recorded (1945, p. 106) "Described from 
five syn-types collected free, from Lake Barrine, 
Queensland, 16 Nov. 1943 (R.V.S.), and a single 
specimen from man, Alherton Tableland, Queensland, 
8 March 1944 (R.V.S.)". 

Reference lo my field and other notes shows that the 
first group ("five syn-types") were my serial ACBI88, 
recorded originally by me as "ACB188 . 5 
larval. ..Running over a log. Lake Barrine shores. Qld. 

With respect to the sixth specimen, stated to have 
been collected (t from man, Atherton Tableland, 
8 March 1944" this is my specimen ACB210A. This 
was the sole specimen of Odontacarus among a batch 
of mites which included many larval trombiculids 
(ACB2I0B, C, D, etc.) and two small (larval) ticks 
(ACC358, ACC359), collected parasitic on ihrce 
soldiers at Gilbey Creek, near Wondecla, Ailterton 
Tableland, Queensland (Map Rcf. Herberton (1:63360) 
392023), on 8 March 1944. The actual collector of the 
specimens was Cpl. E. Grinham, A Company, 2nd/7th 
Australian Field Ambulance, as I was away in another 
area at the time. The other larval trombiculid mites 
collected were later identified as Eutrornbicuia hirsti 
(Sambon, 1927). 

Remarks on the Subgeneric Classification of the 
Larvae of Odontacarus Swing 

The genus Odontacarus (as well as a number of 
others of the family Trombiculidae) was revised and 
divided by Vercammen-Grandjean (1968). Definitions 
were submitted not in the usual terminology of 
acarological or even trombiculid description, but in 
Vereamrnen-Grandjearfs own code. Such extensive use 
of codes in Ihc descriptions and definitions ol 
trombiculid miles has in fact been subject to criticism 
(see Brennan and Go IT, 1977). 

The following is an account of the revision of Odon- 
tacarus made by Vercammen-Grandjean (1968, pp. 


Odontacarus Hwing, 1929 

Ql/Trombkula dentata 

Ewing, 1925 
Leeuwenhoekiini of 
medium to large sue. 

lp- 700-1400 

Remarks and interpretation 
type species, "genotype" 

A new tribe introduced by 
Vercammen-Grandjean on the 
same page (p. 120); on p. I IV 
lie retained the family Leeu- 
wenhoekiidae Womersley, 
1945, and subfamily Leeuwen- 
boekiinac Womcrslcy, 1944. 
U> Index pedtbus, ihc sum of 
t he lengths of legs 1,11 and 111. 
Despite its use by Vercammen- 
Grandjean, it must be con- 
sidered of little diagnostic 
value; e.g. Acomatacarus 
(p. 122) in staled to have an lp 
of 800-1020 (jim). 




Component I 

Component 2 

Component 3 

Trombelki alpha 



ASH, !., A P, Hill 
AW, PSB, LB, Til, 
Gen, Till, Gdlt. Tilll, 
-ASB, -LA, -LN, (A-P) 
AW, PW, ASB, L, LA, 
LB, LN, W, A P, 
AMR, C.cL Til, Till 
AW, PW, ASB, L, I N, 
W, AL, PL, AMD, 
MDS, Gel, Gclll, Till I 

AW, PW, PSB. AM. Gel 
PDS, -<A-P) 

AW, Til, Till, GellL 
Tilll, LA. LN, <A-P>, 

Al. -AMB. -MBS, -PDS 
PSB. L. LH. AM, LA, 


DS, 111 

Note: ASB-ASBa in Souihout <P>X6»i>; PSB ASBp likcwi*Ui 



FIG. 7. A, B Odontacarus southcotti (Womersley), holotype larva; A Dorsal scutum, B Tip of mouthparts, dorsal view. C O. langani 
(Southcott), holotype larva, chelicerae, dislocated as on slide. D O, mathewi (Southcott), holotype larva, tip of mouthparts, dorsal 
view. E O. cooki (Southcott), holotype larva, tip of mouthparts, dorsal view. All to scale shown. (Note: in each case the distal half 
of the cheliceral fang is shown in transparency, so that the denticles on both the lateral and medial aspects are shown.) 


kf.C, S. AUST. MUS. 19(12): 1*9.200 

June, 19H6 

Scutum roughly penta- 
gonal with a nasus, 2 AM, 
2 AL and 2 PL. 

Rye lenses, 2p. 


and palpotibfal claw, 


Subtcrminala or parasub- 
tcrminala more often 
branched or absent than 

Always at Icasl one masti- 
tarsala or a solenidion (s) 
on leg 3. Stigmatae (sic) 
and tracheae present (S+T). 

Ch - 


never ■. 

This means there are I wo 
anteromedian setae, two antero- 
lutcrais, and I wp posterolaterals 
to the scutum. 

The term u 2p" is not defined, 
but presumably means that 
two eyes are present on each 
side, each with a lens. 
This is the code for the <l palpo- 
tarsal pilous formula", 
meaning here that there are 
>even branched setae (plus a 
solenoidala, stated lo be in- 
variably present) on ihe palpal 

This means there arc 2-4 
prongs to the palpal tibial claw 
Those ate specialized setae 
of the pedal tarsi. As accord- 
ing to Vercammen-Grandjean '$ 
statement they may be &b&eM 
or present, and if present, then 
branched or nude, they can 
clearly be allotted no diag- 
nostic significance 
Vercammen-Grandjean, in ihe 
same work, also uses S and T 
with other meanings. 

The code Ch refers to the cheli 
eetae. The notation Jj etc., is 
not explained, but presumably 
refers to the presence ot ftb- 
sence or dorsal (d| or ventral 
(v) teeth on the cheliceral fang. 
The second i is presumably 
an error for £ . as both dorsal 
and ventral denticles are often 
present to the fang in Qdonta- 

Vercammen-Grandjean (1968, p. 120) then proceeded 
to divide Odontacarus into Odontacants s. str., and two 
new subgenera, Taralacarus -and Leogonius. He offered 
no definition for Odontacams s. str., commenting "The 
subgenus Odontacarus s. str. is of no concern in this 
study", doubtless on the (implied) grounds that it whs 
not represented in the "Far East" (a term which was 
used to include Australia, Malaysia, and eastern Asia 
more generally). Odontacarus s. str. was also omitted 
from the diagnostic key given on p. 53 of the same- 
work. Reference to that key shows that Tarsalacarus 
is separated by being a section of Odontacarus sens, 
lat. in which leg tarsus 111 has a dorsal solenidion 
(solenoidala), while in Leogonius tarsus III laeks such 
a seta "but there is 1 mastitarsala and 2 mastitibialae" 
to leg 111 of the larva. Additional characters are listed 


for Tarsulacatusmd Leogonius by Vercammen-Grand- 
jean in that work (pp. 53, 121). Perusal of the character 
listed shows that many of these are shared in common, 
or arc overlapping, e.g. "Odontacarus of median (sic) 
size" as against "Odontacams of medium to large size", 
or "Gr 2-4" (i.e. 2-4 claws to palpal tibia). The only 
characters listed by Vercammen-Grandjean as being 
distinct between the two subgenera, apart from that 
listed above, is Ch-O/v for Tarsalacarus, and Ch=oVO 
for Ijeogonius, i.e. Tarsalacarus has ventral cheliceral 
denticles but lacks dorsal ones, while the reverse 
situation is stated to apply in Leogonius. Onychotrichs 
(cilia lo the pedotarsal claws) are not mentioned. 

The subgeneric classification has been clarified and 
corrected by Goff and Loomis (1977), who defined 
Odontacarus s. str, as "Odontacarus larvae lacking 
tarsala III [i.e. a solenoidala]; mastitarsala 111 present 
or absent", and Tbrsalacarus as "Odontacarus larvae 
with tarsala III; mastitarsala III absent; onychotricues 
absent" Leogonius was redefined by Goff (1979a), the 
diagnosis including: on both claws and 
empodia; cheliceral blade with dorsal and ventral teeth; 
2-3 mastitibialae III present, with basal barbs; masti- 
tarsala III present, with basal barbs. 

By the above definitions therefore, the species (as 
larvae) considered in the present paper come within 
Leogonius, as in fact all Australian, New Guinea and 
South-East Asian species have been placed by previous 
authors, since the subgeneric division Of Vercammeiv 

The characters of the subgenera of Odontacams are 
shown in the following Table 13. 

Classification of Adults and Nymphs of Odontacarus 

As this is the first description of an adult 
Odontacarus, confirmed by experimental rearing, it is 
appropriate here to examine the status of other adults 
that have been placed in this genus on the basis of 
morphological similarity to nymphs reared from larvae. 
So far seven nymphs have been allotted to Odontacarus 
by such experimental rearing, these being: 

(1) O. arizonensis (Ewing), from North America, 
reared by Crossley (I960, p. 196). This species has since 
been replaced in Acotnatacarus s. str by Vercammen- 
Grandjean (1968). 

(2) O, plumosus (Greenberg, iy51). This birtomen 
was used by Brennan and Jones (1959), Loomis and 
Crossley (1963) and Nadchatram (1963). 

A separate genus name, Xenacarus Green berg, 1951 
(since replaced by Xenodontacarus by Loomis and 




Mastitibialae ill 

Xtaiutanafa* tit 

Solenotursuia lit 


Cheliceral denticles 



(Ma Tatth 


















Golf, 1973, on gtounds of pre-occupation) has been 
allotted to this species which was reared to a nymph 
by Cross ley (1960, p. 197). Since Loomis and Goff 
(1973), as well as Vercammen-Grandjean (1973a) confer 
full generic status on tJris and related species, we may 
exclude II nTOOI immediate consideration. 

There arc thus five species of trombieulid mnes 
which have been reared experimentally from the larval 
to the nymphal stage, which may be placed in the genus 
Odontacarus in the sense of Goff (1979a, b). These are: 

(3) O. (Leogonius) australiensis (Hirst, 1925) from 
Australia and New Guinea, reared by C. B. Philip (in 
Womcrslcy, 1945) from New Guinea material, and by 
Domrow (1956) from Australian material. 

(4) O. (LJ adelauleae (Woraersley, 1944), from 
smiiheastern Australia, reared by thi? present author 

(5) Q (LJ audyi (Kadfurd, 1946), from India, 
Thailand and MaJaya, reared from Malayan material 
by Nadchatram (1963). 

(6) O. (I. J novaguinea (Womcrslcy, 1944) which was 
reared from larvae by G. M. Kohls in New Guinea in 
Apiil, 1944 (in Womersley, 1945). 

(7) O. (LJ InnxifH's (Womersley, 1945), which was 
reared from larvae taken from Podargus sp. by G. M. 
Kohls, Dobadura area Of Papua New Guinea, 25 July 
"1940" (probably 1944); recorded m Womersley (1945). 
(This species was synonymi/ed with Q (LJ novaguinea 
by Goff (1979a, p. 149). Nevertheless the nymphs w-ere 
slated by Womersley DO differ significantly from those 
of the Iatler species, and this was not resolved by Goff 
(lo<\ at). \\ ^s therefore proposed to retain these two 
species as separate at present.) 

Womersley thus in 1945 had sound grounds to define 
rhc post-larval characters of Odontacatus (as 
Ac6m£ffHCWVS) t <>nd W allot (u it several speeics of 
adult mites whose previous taxonomic placings had 
been largely conjectural^ these being, as Womersley 
stared (1945, p. 98) "...Rhytwholophus attolus Banks 
1916, R. retentus Banks 1916, both considered 
[Womersley] (1934) as Microtrombidium* and 
Dntrneothrombium dromus Worn. 1939, the last now 
being shown to be really two species" tWomersJey, 1945, 
p. IU9, erected Acomotacarus patrius to accommodate 
this second species). 

Ii is very curious that, despite the widespread 
distribution of Odontacarus (j>JL, to include also 
Acoma(acarus) in North America, Asia and Australia, 
adults of fins genus have been recorded only from the 
Australian continent. This would suggesi, infer alia 
twhen combined with the small amount of information 
OA their ecology!, that the adults live obscure and 
presumably subterranean lives. Thus the adults of O. 
swam recorded here were found in soil in rain-forest^ 
and the original specimen of O patrius (Worn., 1945) 
was collected under h stone, in malice country, about 
7 miles west of Murray Pjridge, SoUtfi Australia, on 
25.V.I938 by the present author, and was recorded as 

'"white in life" which would suggest that that irista*- at 
least is spent in darkness. Four species, O. attolus 
(Banks, 1916), O, dromus (Womersley, 1939), O. 
retentus (Banks. 1916) and O, patnas have 1 been 
recorded as being associated with ants, and in fact these 
arc the only records at capture of these species apart 
from the above-mentioned record for O. patrius. 

All known species of larvae in the genus have 2+2 
eyes, which is consistent with their mode of life as 
ectoparasites upon terrestrial vertebrates. Goff (1979a) 
considers that the New Guinea larvae are principally 
ectoparasites of birds, although elsewhere there are 
records of these larval mites from mammals and 
lizards. In the case of the adults, eyes arc stated to be 
absent in O, swam (see earlier), O. dromus (sec 
Womersley, 1945, p. 110), and O, patrius (see 
Womersley, /or. ttfcj. 

In the case of O. attolus, Banks (1916.. p. 225) staled 
that this species had 1 + 1 eyes, but Womersley (1945, 
p. 108) stated that they "are not now visible"; he made 
a similar statement in 3934 after examining freshly 
mounted specimens. Careful reexamination of the type 
material using phase microscopy techniques (not 
available to Womersley) also fails to reveal evidence of 
eyes, nor was the writeT able to see them in October, 
1946, when the mite bad been remounted by Womersley 
ai some indeterminate time beforehand H may there- 
fore be concluded that this species lacks eyes. 

The only adult species for which previous authors 
have agreed on the presence of eyes is O. retentus 
(Banks, J9I6), for which Banks (1916, p. 116) recorded 
"one eye each side", whereas Womersley (1945, p. 106) 
recorded "Byes present, 2+ 2, small, on distinct ocular 
shields...". Re-examination of the type material 
confirms Womersley 's description. 

This species differs also in having idiosomal setae 
which are terminally forked or branched. Its taxonomic 
placing has been even more uncertain than that of the 
other species mentioned by Womersley (see above). In 
fact Womersley placed it in Microtrombidium 
(subgenus Enemothrombium) in 1934, and in 
Calothrombium in 1937. 

For this species, 1 therefore erect a genus 
Scopiirombium nov, (see further below) within the 
Trombiculidac, subfamily Leeuwenhoekiinae, 

Genus Scopitrombiam nov. 

lype species (original designation) Rhynchoiophus 
retentus Banks. 1916. 

Definition; Adult of normal irombidioid shape, the 
idiosoma not constricted. Eyes 24 2, placed behind level 
of middle of crista. Crista broad posteriorly, narrowmg 
anteriorly to end in a broad arrow shaped expansion, 
the sagitta, provided with two normal setae. Crista with 
a pair of sensillary setae, filiform, at its posterior end. 
Palp slender. Palpal iib»u wjth strong spine with several 
adjacent thickened setae, Dorsal body setae eiliate, 


REC. S. AUST. MUS. 19(12): 169-200 

June, 1986 

terminally forked or branched. Legs rather long, longer 
than idiosoma. 

Remarks: This species is known only from the type 
species. It is a typical leeuwenhoekiine post-larva! mite, 
with a well-defined sagitta to the crista. As indicated 
in the key below, it comes nearest to Hannemania, but 
is easily distinguished by the character of the dorsal 
setae. Its larval stage is unknown. 

Derivation of name: The name is derived from 
scopi-, the root of scopae (f. pi.), a broom or besom, 

an allusion to the branched and ciliate ends of the 
dorsal idiosoma! setae, and trombi-, the root name of 
the family Trombidiidae. 

Scopitrombium retention (Banks, 1916) 
(Pigs 8A-C, 9A-F, 10) 

Rhyncholophus retentus Banks, 1916, p. 225. 
Microtrombidium (Enemthrombium — lapsus for 



If K\ Ot-*»M>r\ 

FIG. 8. Scopitrombium refenfum (Hanks), aduli, lectotype. A Slide-mounted specimen seen in transparency. B Batch of posterior dorsal 

idiosomal setae. C Anus. (All to nearest scale.) 



FIG. 9. Scopitrombium retentum (Banks). Adult. A Crista, eyes and adjacent part of dorsum. B Crista and sagitta in lateral view. C 
External genitalia. D Left palp, medial aspect. E Same, lateral aspect. F Chelicerae. (A, C-F from lectotype; B from ACB748). (All 
[0 scale shown.) 

Enemothromhium) retentus Womersley, 1934, p. 193. Remarks: This species was described by Banks (1916) 

Calothrombium retentus Womersley, 1937, p. 85. and Womersley (1934, 1945). It was recorded originally 

Acomatacarus retentus Womersley, 1945, p. 106. from three localities in Victoria, in each case with a 

Gunther, 1952, p. 41. species of ant. 

M'wrotrombidium retentum Thor and Willmann, Hirst (1928b) was the first to point out that this was 

1947, p. 370. a trombidiid (sens, lat.) mite, not an erythraaeid one 


REC. S. AUST. MUS. 19<U); \&* 20(] 

.fanflj fttoft 

as originally described by Hanks, and he assigned it 
to Microtrombidium without formally giving it a 
binonien. He commented (p. 1027) "This species is 
closely reJated to M. batringunense [Hirst, 1928a]> bui 
the tarsus oi the first leg is considerably shorter ... It 
is closely allied to M. affine (Hirst, I928b| but has a 
shori but distinct nasal process", 

Of the three specimens, each labelled "Cotype", 
specimen ACB750 is the most suitahlc for further 
description, and is therefore designated the lectoiypc 
(see tutther below), 

Redescription of Lectotype, Specimen ACB7S0 
(Supplemented by the Two Paratopes) 

The specimen in its present state is a squashed mass 
mounted on a 75-25 mm slide in a transparent, 
yellowish water-misciblc medium, based on eithet gum 
chloral or polyvinyl alcohol, with picric acid used as 
a stain (Womersley favoured the use of picric-acid- 
containing media in the later 1940-I950s and possibly 
later). The squashed mass is the remains of the 
idiosoma, and is about 2.7 mm long by about 2.25 mm 
wide, indicating a considerably smaller original 
specimen (Banks recorded the specimen(s) he described 
as 1.3-1.5 mm long and "about one and a half times 
as long as broad" i.e. ca 0.9-1.0 mm broad. 

Crista is a rod set in a narrow tapering scutum, 
widest posteriorly, and ending anteriorly m a broad 
sagttta; crista and sagitta with combined length of 306. 
Sagitta bell- or probability-curve-shaped anteriorly, 
borders smooth, posterior margin reflex-angled; sagitta 
82 jtm across by 52 deep. The corners of the sagitta 
are sharp (i.e. lateral angles). Sagitta bears two seobalae 
which are both broken off, at 21, 25 length; they ate 
not broadened, but appear normal and almost smooth, 
From specimen ACB748 these can be seen to be tapered 
and pointed, 68 long. Sensilla bases normaK at 
posterior end of crista, set in a sensillary boss, centres 
37 apart; sensillary setae filiform, about 180 long. 

The crista and sensillary boss are invested by a 
narrow scutum, which tapers anteriorly, leaving the 
anterior third of the crista without this. The scutum 
is free oi setae, but is set in the normal plicate idiosomal 
skin with its characteristic setae. Behind the sensillary 

boss is a conical continuation of the crista, probably 
subcuticular, about 50 long. 

Eyes 2 h2, each lateral pair set in a small oval plale 
about }& long by 25 across. Anterior eye considerably 
the larger. Cornea 18*20, posterior eye more or less 
transverse, with cornea 16x9. Eyeshields set at the level 
of the junction of the middle and posterior thirds o\ 
the crista; centre of each eye shield about 171 distant 
from the midline of the crista. 

Various dimensions of the type and paratypes are 
shown in the following Table 14. 

Dorsal idiosomal setae arise from a small papilla set 
in a broad plate. They are heavily dliatc, more so 
distally, and vary considerably in length. A clear 
division between two distinct length-groups cannot be 
made. The setae tend to split or branch in about their 
distal third, and these branches may branch again. As 
all parts of the seobillum are heavily ciliatc with coarse 
cilia, the end of the more branched setae appears 
commonly besom-like, these remarks applying parti- 
cularly to the longer setae. In the anterior part of ihe 
dorsum there are comparatively few of the longer setae, 
except immediately around the sensillary area of the 
crista, but they arc more numerous towards the 
posterior part of the dorsum of the idiosoma. 

Ventral surface (partly from ACB749) provided with 
numerous but shorter ciliatc setae, more uniform, and 
with only a slight tendency to branch terminally 

Genitalia oval, 235 lonu; by 165 wide* with three oval 
suckers alongside the imroitus on each side, anterior 
sucker 57 long by 29 wide, middle 48*34, posterior 
49x36. Each valve of the external genitalia with a single 
row of tapering setae 32-45 long. No internal 
chtiinization seen to genitalia, so the specimen ACB749 
is possibly a female (damage to the specimen making 
this a little uncertain). 

Anus oval, about 62 long by 51 wide, with two valves, 
each valve with 4-6 ciliate setae 31-35 long. 

Legs moderately long and thin, appearing normal 
tor a trombidioid mite. I 1670 long, II 1270, III 
incomplete, IV ca 1820 (lengths including coxae and 
claws). Legs with a normal degree of setatlon. Many 
of the ordinary setae (seobalae) are unilaterally and 
regularly filiate, thus making them classifiable as 
pectinalae (sec above). Dimensions of leg segments o\' 




iiificuiion Number 





r -' 











ca 2K5 

ca jhw 



sag ill a) 






362 - |28 

Ml 105 

1«1 X 'JO 

3MI lid 














ca ux) • y-' 

278 - 95 


















IK >1bl 

2-.«. 5m 

:'■'. •'< 

* This leg has been dislocated and damaged, but I presume thai il tu h-.-t-ii cdirccftv identified 
r Damaged specimen, and only a few of ihe lonyer *et.Ht jrc available 



leeiotype: tarsus I 362 long by 128 high, tibia I 339, 
melius I 24S, tarsus [I 252 - 70, tibia II 245, genu II 168, 
tarsus til 265*86, lihia III 286, genu III 195, tarsus 
IV 300-97, Ohio IV 402, genu IV 274 (all tarsi 
measured without claws). (The preceding assumes \ 
have idenl i Tied legs HI and IV correctly.) 

Tarsal claws sickle-shaped, uncinate, sligblly blunted, 
appearing of normal irombidioid fades. 

Palpi rather slender, normally setose, Palpal tibial 
claw stout, somewhat blunted. A blunted large dorso- 
media! accessory claw (a thickened seta) alongside claw, 
and several smaller thickened peg-like setae behind it. 
One long whip-like seta present on lateral aspect of 
rihia terminally, over-reaching claw. Palpal tarsus an 
elongate oval, wilh usual selation of ciliate and sensory 

Chcliceral fangs slendei with about 12 indistinct 
teeth along the flexor (upper) border. 

Material Examined 

I k only material available is the three individually 
mounted specimens of the original type series. 
Womerslcy stated (1934, p. 193) "As the specimens have 
been remounted for further examination . " but 
only two of the specimens have been remounted from 
the original balsam lo water-misciblc medium or 
media. 1 suspect thai the two slides remounted in the 
period 1933*1934 were remounted again in about 
1944-1945, for reason.'- given below; Details of the three 
specimens are as follows: 

(I) Specimen ACB750. Leetolypc. Specimen 
mounted on a slide 7.*S mmv25 mm, squashed, in 
yellow-stained medium, The only trace of what was 
possibly the original labelling is a small pink label with 
"Cotypc" in red (LHS). On Ihe RHS of the slide is a 
label m Womersley's writing which reads Caiathrom- 
trium/retentus(Bmks)/CoA [m redj/with Indomyrmex 
mtidus/Ocean Grove, Vic/Nu. 17 Lea Coll. 

• 2) Specimen ACB749. A slide similar lo the 
preceding, with the mite squashed and disrupted into 
three main parts and various lesser parts, in the same 
medium, ll is labelled wilh a small pink label "Cotypc" 
as in the preceding (LHS); on the ftHS is a label in 
Womcrsley's writing Calothromh/um/retentus (Bks)/ 
Co- 1 [in red) Ocean Grove, Vic/No. 17. Lea Coll. 

This specimen is clearly a female, as about 20 
spherical or spheroidal eggs can be eounied in trie 
fragments. These have a normally pigmented chorion, 
jjld measure about 140495 by J20-I60 in the least 
squashed Specimens. There is no evidence of an embryo 
in any oi' the eggs. These eggs appear as normal 
Prostigmata-typc eggs. 

AH legs arc disrupted, and the lower (separated) part 
of the idiosoma retains only the trochanter 1 and 
trochanter II on one side, and trochanter IV on the 

This slide contains fine acicular crystals which would 
confirm ihat a is mounted in a chloral hydrate- 

containing medium, at least in part. At the time when 
Womersley remounted it in 1933-1934. tie was usinjt 
exclusively gum-chloral media. However, a laici 
possible use of a polyvinyl alcohol medium cannot be 
excluded, there is clear evidence of two zones in the 
mountain in the slide, The slide contains also some 
instar of an insect, possibly a primitive one, and also 
a rather stout and short trornbidioid leg, different 
character from the legs of Scop'Urombium rrfenrum. 
How this extra biological material got into the 
mountant is purely a matter for speculation. 

In my opinion the specimen is in no condition to be 
designated a lectolype. 

(3) ACB748, A slide of an almost undamaged adult 
mite, mounted on its side in balsam (thus Indicating 
that Ihe other two specimens have been remounted 
from balsam to watet-miscible media), with a cover- 
glass of 19 mm diameter, l( bears in the writing believed 
to be that of Nathan Banks (Dr E. G. Matthews, Senior 
Curator of Insects, South Australian Museum, who is 
familiar with the writing and labelling of A. M. Lea, 
advises that this writing is not Lea's, so it may therefore 
confidently be attributed to Banks.): <R.H. label) Rhyn- 
cholophus/retentus/Bks/Microthrombium [this word 
in Womersley's writing] /Ocean Grove/Victoria/ with 
Indomyrmex nitidus/Corype/No. (I interpret this 
figure as 14) 14/ Lea coll.; ( L.H. label) (in Womerslcy** 
wntine): Calothrombium retentus (Bks)ZCo-t |in red] 
/Ocean Grove/Vic No. 17/Lea Coll. 

As ir ts mounted on its side with its legs partly flexed 
(sec Tig. 10), it is clearly not the specimen mainly used 
by Banks for his original description. Despite the RfcC] 
that it is the only specimen in which the original data 
remain i\u the slide, it is not the most suitable to be 
•ed as the lectotyr>< 

Further Remarks on the Type Series an/1 
Leeiotype Designation 

Banks (1916, p. 226) recorded three localities lor this 
species: Lai Lai, Ocean Grove and Sea Lake in Victoria. 
Despite this, all tluee slides are labelled "Ocean Grov^, 
Vic." and "No. 17 Lea coll/' in Womersley's writing (set 
above). There were clearly at least two original slide* 
(and probably three). The remaining balsam mount 
(ACB748) has clearly never been interfered with. Tbe 
balsam mountain has yellowed in its more peripheral 

According to Banks, the specimen Irom Lai Lai \ 
captured with tiie ants of Polyrachis hpAacunrha, while 
the specimens from Sea Lake and Ocean Grove were 
captured with Inrlomyrmex nnuius. Accepting ihe 
accuracy o\' the data on slide ACB74S in BankVs 
writing, and if we accept that the notation o\' 
Wornersley on slide ACB750 about the species being 
"with indomyrmex n nidus" h eorrecL it may be 
deduced that rhis specimen, tbclectotype. in lad came 
from Sea I^akc, and that specimen ACB749 in fact came 
from Lai Lai. Whether we arc entitled to place any 


REC. S. AUST. MUS. 19(12): 169-200 

June, 1986 





FIG. 10. Scopitrombium retentum (Banks). Adult, specimen ACB748, mounted on its side. 



reliance upon such a deduction is conjectural, in view 
of Womerslcy's other obvious inaccuracies with data 
on the slide labels. 

All three specimens have the distinctive type of 
dorsal sctation recorded, and the tabular data of the 
ehtrinucd parts also confirm that these three specimens 
arc conspecifie, in addition to the opinions of Banks 
and Womersky. For re-description, a lectotype needs 
to be selected. For this purpose, all three specimens in 
their present state have some defects. Specimen 
ACB748 is the only one retaining a presumably original 
label, bur is unfortunately mounted on its side. 
Specimens ACB749 and 750 have been badly damaged 
by being Temoumed at some stage, undoubtedly by 
Womersky, and possibly at the time of his 1945 study. 
Despite the serious damage these latter specimens arc- 
more suitable for the usual standard descriptions used 
in prostigmatic and other mites, as the crista, eyes, 
dorsal setae and other parts remain deseribable. 

Unfortunately Womersley was afflicted by poor 
vision, and he was often extremely destructive to 
specimens in an effort to make them easier to see. He 
likewise erred frequently in transcribing locality and 
host information and often destroyed earlier labels. 

One may therefore conclude that Womersley mis- 
transcribed the locality and other data for the two 
specimens ACB749 and 750, and presumably one of 
these came from Lai Lai, and the other from Sea Lake, 
hut it appears there is now no way of determining 
which came from which. 

In his writings upon this species, Womersley even- 
tually (1945, p. 107) was aware of the three different 
localities in Victoria from which this species was 
collected, but this is unfortunately not reflected in His 
slide notations, which were presumably made first for 
his 1933-1934 study, but then re-transcribed for his 1937 
study, in which he used the generic name Calothrom- 
bium Bcrlcsc, 1918. 

There is nothing in the description and figures of 
Banks which allows the selection of one specimen as 
against the other for designation as a lectotype. Several 
important diagnostic characters were overlooked or 
misinterpreted by Banks. Thus he placed the species 
in the Hrythraeidac, instead of the Trombidiidae tas 
then understood), He recorded only one eye on each 
side instead of I wo, and Tailed to see the sagitta to the 
crista. He gave almost no morphometrie data, and his 
description of the species is such thar> as far as present 
evidence allows, it could equally well be applied to 
either of these two specimens (or even ibtaKX as was 
probably intended by Banks in any case. 

II is regrettable that Womersley re-mounted these 
specimens so crudely, and mistranscribed at least some 
of the data on the labels. 

Ol the choices available, I select ACB750 as the most 
suitable for redemption, and hereby designate it rhc 

Taxonotmc Placing o/Scopjtrombium retentum 

This species has had a rather chequered history of 
taxonomic placement. Originally it was placed by 
Banks in the family Erythraeidae. Correctly transferred 
to the Trombidiidae (s.I.) by Womersley in 1934, it was 
Initially placed in Microtrombidium Haller, 1882, in 
the subfamily Mierotrombidiinac, and later (1937) 
transferred to the genus Calo/hrombium in the 
Tanaupodinae. In 1945 Womersley recognized its 
affimties with other Leeuwenhoekihae, and transferred 
k to this subfamily, which he elevated to a family, its 
a member of Acomutucums Ewing, 1942. 

The suite of characters listed above, however, set this. 
species apart from the other members of the subfamily 
Leeuwenhoekhnae, and distinguish it from the Neo- 
trombidiinae Fcidcr, 1955, whose systematic position 
was clarified by Lindquist and Vercammen-Grandjean 

The following is a key to the adults and nymphs of 
the subfamilies o\' the TrombicuJidae (after Crosslcy, 
I960; Lindquist and Vercammen-Grandjean, 1971) for 
which adults or nymphs are known. 

I. Sagitta of the crista with one seta or without setae; >T 
such a sagittal seta is present it is usually expanded 
somewhat or well ciliatc. Palpal femur with few setae, 
equally numerous as, or less numerous than, in palpal 
genu, Sensillary setae of crista eiiiate (except Blankaariia 

Oudemans, 1911) Trumbieulitiae 

Sagitta of the crista with two setae. Palpal femur with 
numerous setae, more numerous than in genu. 

Suisillary setae of crista filiform 2 

2(1) Lyes 2*2. Body setae conspicuously triramous, Coxa 
of al least die posterior leg with mosaic ornamentation. 
Two pairs of elon«a(e elliptical discs or suckers to 

external genitalia in botn nymph and adult 

. . . , i . Neolrombidiinae 

Eyes -♦ 0, I t i, o\ 1 + 2. Body setae ciiiate, and may 
he branched or ol other form, hut not conspicuously 
triramous. Leg coxae without mosaic ornamental ion. 
Adult with three pairs of oval discs or muckers along 
each side of external genitalia Leeuwenhoekhnae 

Generic Classification of Adults and Nymphs 
in (he Subfamily Leeuwenhoekitnae 

Recent classifications of the status of the various 
genera and subgenera ol a number of the Trombiculi- 
dae, particularly the subfamily Lecuwenhoekiinae* 
together with the present study of the adult of OdonJa- 
carus, and the separation of Scopitrombium* allow 
some revision of the generic classification. 

The following is an attempt at a key to separate the 
genera, for the adults and nymphs, of the Leeuwcn- 

! . Eves absent 2 

Eyes present — 4 

2(1) Posterior dorsal body scue in two distinct forms, rhe 
shorter spindle-shaped, pointed, ciliate, the longer 
sword-like, with its proximal part ciHatc .... 

Odantavants Ewing ]92V 

PoMenor and other body setae lend to be locally uniform 


Rfet & ADM. Ml V 19(12): I69.2UI 

June, 19M 

3(2) Posterior dorsal setae leal'like, \x$s i»oi longer than 
hody. Tardus I pytiform. Crwlkcral blades narrowed, 
neai -|> sheai like ,, . M'/nir/wi/i/ fcwing, 1944 

Posterior dorsal setae somewhat expanded, but not 
IcafliWc. CnehLtral blades shearlikc, not narrowed 
. . .. AcvnmfacoTus fcwing, 1942 

4(1) Eyes I . I ... 5 

Eve-, 2. 2 . - 7 

5(4> Sagittal SCtSe expanded. Cheliecral blades long and 

dagger- like Xenndontaivnts Loom in and Gofi, 5 *>7 3 

Sagittal Nctae W\ expanded. Cheheeral blades not long 

and duggei-likc- . 6 

*pi Posterior dorsal setae not expanded, eifiatc, and ending 
in long attenuated lips. Crista promintnUlv puuetale. 
Lees longer than body Tarsus I cylindrical, aboui 2.5 
times at tofljj as high ....... Chaiia Brcnnaa, 1946 

Poster tor dorsal setae somewhat expanded, nnt ending 
in long attenuated tips. Crista a narrow rod, not 
prominently punctate. Legs of normal length. Tarsus 
I eyliudiieal. about 6.2 limes its long as high . . . 
Comatucan-ts fcwing, 1942 

7(4) Body constricted, to a ligufe-o! -eieht shape. Posterior 
dorsal settee not expanded, rather elongate and ending 

in a hook-like process Tarsus 1 cylindrical 

, Hannemania Gudcmans, Nil. 
Body not constricted Posterior dorsal setae terminally 
forked or branched, farsus 1 an elongate oval , 
Scapitrvmbium gen. run 

in preparing the above tabular key I have accepted 
the separation of the Neotrombidiinae (or Neotrom- 
bidiidac) from the Leeuwenhoekijnac, proposed by 
LindquLst and Vercarnmen-Grandjean (1971). 

I have also omitted the genus Parvithrambrium 
Andre, 1962 (whose sole species is R crassirarsate 
Andre, fS62» from Angola), which its author placed 
in the Leeuwenhoekiidae. In the somewhat speculative 
revisions of the classification of the Trornbtdtoidea 
which have been proposed in recent years (Vercarnmen- 
Grandjean 1973a, b; Feider 1979) the genus Airvr- 
thmmbtum has tseen placed in the Anomalothrom- 
biinae Feider, 1955 (later elevated lo Anomaiothrora- 
bidiidaeand Aoomalothrombidioidea by Feider. 1979) 
along with Anomaloihrombium Andre", 1936. 

Anomalothrombium adults, the only instar known. 
possess a small ,k naso" with two setae somewhat 
resembling the sagitta of the Leewenhoekiidae and the 
Trombtculidae. This genus appears to resemble the 
genus Spelaeolhrombiurt? Willmann, 1940, and as far 
as descriptions allow (Andre, \936 § 193S, 1939, 1*45, 
1958: Meyer and Ryke, 1960 (for A. curiosetosum 
(MeyeT and Ryke)), Robaux, 1965) could be placed in 
the subfamily Spelaeothrombiinae as redefined by 
Robaux (1968, J 972"). 

It appears probable that The Interrelationships 
between these mites, at present m a srate at tgxonomic 
instability, will be clarified only when the larva-adult 
correlations are known, at least in the first instance. 


f am greatly indebted to the South Australian 
Museum for access to type and other material in the 
acarological collection. I also lhank Mr L. G Veitch, 

Principal Research Scientist, CSIRO Division ol 
Mathematical Statistics, for advice and asslstaruv ill 
statistical matters. 

The National Health and Medical Research Council. 
Commonwealth of Australia, provided financial 
support for these studies. 

kk»:rkn< ks 

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retted hj £">H and iJQOmJdt, 1974) 
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• .< IKR M i 1979*, lhegcnusOf/onr*/ii/nt>(Acnri: Imnih.ctiliiJac) 

En New t lufiiea wltfi dcsc-ipiious of tour new -.pccies. ./, rnt'd 

t a -1 f t M, I IV? 1 *!.- A new spe. Iff* dl ' toMMCBfttJ I Vari. Troru- 
biculfUae) intcStlfiu ID U fid lodcnt, in Papua New Guinea. /. maJ 
tinlomQl. u> 140 141. 

GOH-. M I I9KI live new species ol rfilfiftCj* |Ac*Ti 

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pv^osotnal pi. iic. 7. /w<7. I'ntotnol 15 SO 5] 
(.tUP, VI I.., iiml lOOMlS, R. U 1974 A new species oj 

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UOII, M. i., LOOM Is. k. It, Slid HRbNNAN, J M. 1972 

(hioiiUnarus-itmnnus (I winv. I g 25) & "he sCllloi sy(ion\m ol 

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■ ■ ■ / ■ ■ 5-481 
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I V.i.iiMi ltombieuh<Ui>) ifrih' I »/tf Puh/lCn. Btttamol. H 

<.mi-N(5hKU, U. I95l> A new •athe.eiius of Acomatacurus From 

Kansas |-V.-a.ii.a ITomD/CQtKtaCJ- I r*ara\\tot. .17: S25-527- 
GKE£NB&RO, 8- 1952. a revit-w oi ihc new woild Acomatecorus 

|-Vr:.n,ij- TnjinbkuliJai?). A$m. &1t. $Ql -I""'/- •*$ 473-491. 
GUWTHfcR,< I.. M, 19.S2. \aicck lisi oi ihcirornbiculid Icuvoe 

ol' Asi;i .tiid Au^ntlJMii. Pr0C 'inn Nor, \.S.IK 77: 1-fSU. 
HALLtF. M_ I0K2 Kciirag /ur Kcrminisb der MittTBnfauim 

Wurnwntiengs. Jahmxh. ler. m\M Naiurkumie'm Huntembw 

3a Z03-33S. 
1IIRSI", S, »y^s. or, a IrarVcM bog y rvuwenhtit'kitt misfrultcnsu*, 

•>p. n.) alU(Ck(rtJ3 Iihmuhi b^lfif. si Nvdm.'v, New Suiilh Wat*;*. 

Trans, ft Sot; TTouyv Afttf d! Wft tt 156-15 J 
HIRST. S. pr^Sa. On Rome new Australian mitC9 v\ lll« tainilies 

rrombidiiJar anJ I - viln.ieii.ljc dfffl; Mfig fftf/. i//-s7. (IBf) I 

HIRST, S. 1928b On some AiisiraliaiiiptxicsoriKimbidiitJae Ptm* 

Aool S,n LoruL Mn 1*7): UUMC04. 
IKMl I I l>Ji... H, P^3J3, AhhIvsim.I ( i complex of siaitstical vnrHtl'k-s 

into principal con.pnncnis J e</iw P'ivchnl 24. 4|7 44I, 

KISH1DA, K 10?^ IhuHidunoii ol Aemtcurus Kishid;i. 19:5. irrd 

ril .v maktrtoi Kbbitlrti I9?5j. A>tsrracM of tire 7 iri Mating oJ 

Ihe Iokvo C-ar.»Mioloei' \il W<eiv. No. 2459: -II Iquuted by H. I s. Rfehioka. I9S7 in U.idid nckfol lapan, Kwa 
mo i ii<- R>n».y" 'sUm-K, Depi I Niamola^ <fl6ih MrJlcal 
General liboatury, * >onp Aimim. i.ipar.. and rhe LT.S. Arm^ 
Medical Reaenrch l-n-ueiHuiin, F»i Eftrl p 'lj 

l AWRtNCI- Ft I 1941): P4tW I LrV&l loims of Sdlfiri Ahiciiu rmrcs 
from anlin.pod A09M " Ifl - JWWff/ Wto 9- 40I-40H. 

i iWftthW I- R I 1949 I he larva! trtmtbli -ui.d n\ifc$ oi South 
i .i i.i ,,., lrt/1. \-vi,// A/;/s. ll 405486, 

I l\|)g»i|Kl t l . I .,and\T.KCAM\H *N-<;RAMJJ» AN, t* tt. W\ 
UeviMoti 31 ilu-i InggCI lifcc larvae "I UltBltfUW ^><>in>nit^tiutu 
1 cn-irdi and MonUftJtUG W hutori. -,v i 1 1 1 a ledchniOdn "' Lite 
.tlbfaillih N:*.huMi|Tidil[i.ic leldtr Ml • lie lTOTnbTiliWw: (Allium: 

rr-^tij-inaiai y amui RnumatogtR I0.V 15574590 
IwnMIS, R H »*J'f» tin- dllgger mites of Kansas lAcann:.. 

Iruinbicui'dac) th>,\ fc)«.flj.< .St (. «»//. 37: tl»35-l4& 
U HiM IS. R Ii, ;nult krtS'-it RY. 12 A t . Jr.. i^V New .peers and 

nevv rtcorifc oi i hlugeM rAarino; iMnihitutidaei Tto&$. 

1- hiMIS R_ K, inuM-i'l | ( M, L 1973% A f*V l ' ; ' 0O» 

ip*cies ol Noilh Amsrifcail Iccuwciihoc^MMccluefc't^t.Varina: 
lininhiciilidae) ,/ fried* f ttiomot. W II 3-7 P 

M.ii'lliH ii. R N 1444 NiVCn on ihe d^iribuium and hahit- 
u lip.i i d iniicy m Queensland and .New viliuica. Mnl J, 

\«:t., IW4 2. 5^3 5*% 

McCfJl "H H KA IVT*i. Studies hiihc o'Uliol .ni >cni»- iv[-'-n. 
A;*7y ./ 4U,« I'M/. I: FlT-738 

M,< I'lIOf II, K.N VHi nicad.c.iHin I military scrub Ljyhio 

CODlrpI to coihan iltusUi |ft •:/ J lUAfi. »°47 I 449*494 

Mt'YtR. M K. R, and RVKH, P. A. ) I960 IrombiJiidac (Acaritta.' 

Ptostiginalal cOUeCCdO fr<»ai Souih Aliican grass and IQU. ./. £/r/. 

Sat S- A/r. >3: 194*3118. 
NAIXHATRAM, M. 196?. 1 he larva and nymph ol" OJant,Hun^ 

audvi {K^dltiro) tAcatina: Trombicididaet. Pacific //iwu.s .V 

535-540 T 
DUDEMANS, A. C I9i(t. Av;.r..i..[Ai-.chcannrci*keningv*n XNXUJ. 

/j/;/ tffv., Arrutmrtam 5: 83-90- 

OUOfMANS, A. L. I9LI. Acarolot'ischeaanieekeninyeii WNV1. 
£W, ^/ \msterdam 5: 1T"'-l- n '. 

OUDBMAN5, a, c t9U. Die bis _it.-i/t brkflxinntH Laiven vom 

TliloiTit'<idii..lje Mini l i ynn,iLidac lir:,ondrrirr HcfiLkHcho 

tune doc fur den ^enaction siMdMcftvu Ancn 2opJ lahrh., 

Abi i, Supp!. MV. N.j L pp I-2V). 
RAOK1RQ, C. S 1946. New species or larval mile* (Achilla injifl Mjinpur Suit, India. PftSC. Zoof. Soi 

London IH.: 24 7 265 
RhFO J 1. 1973. Cvmutacarus, lormerlv .mUl-chus ol 

Leeawenhockiu, rcMorcd 10 yenerie siaiu»«, willi ile^cripnoti -n 

a new mcvic- I Acarina; Trornbicuhdael 1 PTfiflflL Fniunml. 10: 

RCEa J I. Wnj ftRFNNAN, I. \1. W75. The -uhoinily 

i.ccuweiilincVinae (sicl "i I l.c rttaH op.. - (Acarioa: 

rromhieululac). Bnvkom )f>unz Univ. Set. Bull., Biol. Ser. 20: 

I 42. 
ROBALX, V. 1965 Sur queiqaes MicroiKionihidioiae dc Cote 

ITyoirc lAvajirtft, riir».ni!<iditdac;i. fr^VRfl Qc^f. ftw/. -S'^/. 2: 

63 K4. 
ROIiAUX, P 1908. I' ■ihruinbiinac: tc statu! du (y&nrv 

Tvphlnthromhium Bcrleie 1910 ei description de deux noHVCnUl 

teen:, (Acjfi I hr.>mbiJiidae). Acaroloyht Of- 44-64 
KOHAliX. K l" 7 2- In noioeau Spelaeoihrombiiuae v^vfinicolcdc 

Cuba: Sfwothmnibium mottUi lihtd n y, n sp. (Acuri 

Thrcmbidiid.jc,i. Annolcs \pele<>l 11 7H7-791. 
SAMBoN. I W 1927. I he •Viub-itch '* mite of north Queensland, 
i w tpCClM oi" I'mntf-iculu Ann, Muv. nnh Hist l9i 2(1 

SOUTIIvini, R V, I9f>7. Ihc genu., Xcowiitucaru.s OVanna: 

rronibiculidae). I Description ot three new species from Irimrv 

Hay. iionli (Queensland. Irons, tt Soc. S, \ust 80- 146 155. 
SOUIMCOTT, R. V. 146ft RcMSH.m ol iltc ^enuv Ctiurlfiutnu 

Ohdemans (Acarina: Frvtliracidae) AllSi / 25ooA 14 6K7-8W 
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letresirtal Ariliropods". R. V Souiheoli Miiehaiu, Snuih 

so^UiHCOl I. R. V I9?r». Aracltntdisin atul allied syndrom, n 

Mm \n-iulid.i region. RtC Add Chtldr tfosf*. i 97-186 
SOLTHC^irr, K V 197H ' l AiibInlian Harml'ul Arachnids nnil I tu-u 

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.t ril.. I urc. AVc S AUSI Mus V> 145 K-S. 

SOUTHCOn, R V. I98f.b Desenpnonol QflonUuant* \eit<hts V . 
nov. (Acanna Trombitulidaei. Rec. S. Auu Mus. 19: 213 217 

IAYLOR. I H. 1946. Section I. Description, hi "Spiderb. Tick s and 
Mites indudiny. ihe SpeLic-. llirniru! to Man ifl Australia and 
New (Tninea'M Tavloi. I H. and Murray. R L edO. p|> 7 -2U. 
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vicdu.ine. University of Sydney jmi! ComroonwcaiCl pop -9 

THOR, 5.1 :nid WILLMANN. L . 1947 Irouihidiidae [Ms Jifrrifh 

7tb: |K7 541, aiH-^VltvL 
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UMiig metiic dala ifom larval specimens of three species ot 

0(iorthtLurn\ (Aainr.,c. tu.iiibiculidac) 4f*W Ji A>of. J3' 


VF.RC 'AMMLN-ORANIUFAN, P. H. 1956. Ufl Lecuwenhoc^ 
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(Mtffmuruy) hurrtti. it s^., n. sp. (Aeannat Ann. Mu$ ""•' 

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VIKIAMMLNA-RANOTI-AN. P H 1971a. SlU !r- ilaluJ S ." 14 

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IT Re AMMPN i.M \NIJ9LAN, P II. 1973b, las.-noM.Ic Jos N«o 
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June, 1986 

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The early history of the discovery of Speleognathus australis Womersley, 1936, is recorded. In 
Australia over 100 specimens of the mite were observed personally on the water surface of three 
cattle- (and horse-) troughs between 1934 and 1939 at Glen Osmond, South Australia. Despite 
hundreds of observations in the subsequent years on the fauna of cattle-troughs and other water 
surfaces, including the original sites at Glen Osmond, and many other sites in Australia and Papua 
New Guinea, no further specimens of this species have been discovered in Australasia. 




honorary Research Associate, South Australian Museum, North "terrace, Adelaide, South Australia 5000 

(Manuscript accepted 23 January 1985) 


SOUTUCOI I, R v. 1986. History of the discovery oi Speieognathus 
(msfrulis Womersley (Acaritta; Tromhidirotmcs), wilh notes on 
Ms natural history and behaviour. Rec. 5. Aust. Mus. 19(13): 

The early history of the discovery of Speieognathus 
australis Womersley, 1936, is recorded, In Australia over 
100 specimens of the mite were observed personally on 
the water surface of three cattle- <and horse-) troughs 
between 1934 and 1939 at Glen Osmond, South 
Australia. Despite hundreds of observations in the 
subsequent years on the fauna of cattle-troughs and 
other water surfaces, including the original sites at Glen 
Osmond, and many other sites in Australia and Papua 
New Guinea, no further specimens of this species have 
been discovered in Australasia. 

This species has been recorded from two other 
locations as nasal endoparasitcs of bovids, once being 
found in the nasal fossae of Zebu cattle, Bos indtcus 
L.» at Astrida, former Belgian Congo, Africa, and on 
another occasion in the nasal fossae of the North 
American bison. Bison bison athabascae (Rhodes), 
near Cache, Oklahoma, United States. 

Observations of mite behaviour on the water surface 
are also described. 


Speieognathus australis was described in 1936 by 
Herbert Womersley (1889-1962), Entomologist, South 
Australian Museum. In his original account Womersley 
(1936) gave the following field data; 

"Locality— Type, one of two specimens taken in moss 
at Glen Osrnond v Adelaide, South Australia, in July 
1934 (R. V. SouUteoU); four other specimens from 
same locality in January 1935 (R.V.S.)-" 

All of die specimens collected had been caught 
running over the surface of the water in three (or less) 
catrle- (and horse-) troughs, at Glen Osmond, South 
Australia, and this information had been passed on to 
Womersley. At the time I had been supplying him with 
large amounts of moss, leaf litter and soil from various 
localities in South Australia and Victoria, for his 
studies on [he Collemboia and Acarina* In view of the 
volume of this material chat Womersley was handling, 
his initial mistake is understandable. As 1 was only 16 

years old at the time, perhaps understandably he did 
not check the accuracy of the locality record before 

However, this species of mite was unusual, not onlv 
in its morphological similarity to the ercynetid mites 
(Riccarcloella) that are common upon slugs in the 
Adelaide area, but also in its behaviour. Its manner of 
running on the surface of water showed that it was 
quite at home there, As far as 1 was concerned, there 
was never the slightest question that every specimen 
found in Australia up to that time (and subsequently) 
had been collected only upon the water surface of these 
catlle-l roughs. I had found that in collecting mhes, 
Collemboia and other anthropods, the surface of eacn 
of these cattle-troughs (about 45 cm across by about 
150-180 cm long) provided an area for sampling in 
which all specimens were clearly visible, and at the 
collector's mercy apart from the occasional watcr- 
strjder, water beetles, or other truly aquatic insect, and 
could be captured and bottled in alcohol without 

On the surface of the water, these light brown 
Acartna stood out by appearing to be either fully 
aquatic, or at least adapted to a life at the surface of 
the water. Specimens were collected from 1934 onwards 
and taken to Womersley, and repeated observations 
were made upon their behaviour by myself, both in the 
field and in the laboratory. These repeated observations 
were described to Womersley In fact, on one occasion, 
I took Womersley, at his own request, to the troughs 
in question, some time before the publication of his 
1936 paper, and possibly as late as April-May 1936, the 
exact date not being at present available. (Womersley 
and I lived in adjoining suburbs at the time, and the 
troughs were within easy walking distance). 

It was recognized that the miles' unusual mode of 
life at the water surface indicated an adaptation to a 
moist surface, and the possibility of their being 
nasicolous parasites of cattle and horses was 
considered. Following the joint inspection, Womersley 
arranged for the noses of the caltle to be swabbed by 
a veterinarian on 20 May 1936 (see further later), but 
no mites were revealed. 

When Womersley's paper appeared in 3936, I again 
advised him that all specimens had been collected at 
the cattle-troughs and asked him then and on several 


ku s v si MUS i i Mi5>::oi2i: 

June. 1986 

later occasions to correct the mistake at some ttrrifl ill 
the future; This Womersley declined to da 

In I94S Hi/abeth Boyd described a secmd species 
of mite in the family Speleognathidac. which 
Womersley had erected in 1936, and minted it 
Speleognaihus Sturm, since it had been obtained from 
the nasal passages of starlings {Stnrnus vutgaris I ) In 
North America. (A specimen of the mite had also been 
captured from the boat-tailed grackle. CilSsfdl \ 

in 1952 Lawrence described an intranasal riliK fltoO 
a South African toad, Bufo regulars, which lie named 
Riccardoellu eweri Lawrence, 1952, thus recognizing, 
as Boyd (1948) had done, the etvynetid affinities of 
these miles. (This species was l3ter removed 10 
luwrenvurus by Fain in 1957, and is now known as 
Luwrencants eweri (Lawrence, 1952).) 

In 1952 also, Crossley described a species ol 
intranasal mite from a smnl* number of specimens 
obtained from the domestic pigeon, Columbu Ihto 
domestiva % in Texas, U.S.A., as Speteognothu\ yfrwwy. 
That species is at present known asOpiuhaloiox'nu'fr- 
strioms (Crossley, 1952) (see Fain 196), [)oi | 
1969). Curiously enough, (be discussion in CrossIevN 
paper (1952, p. i86) contains the following; 

"The type species of the family Spclcognathidae, 
Spetepgn&thUS australis, was found in moss and has 
never been reported from a bird. In 1948, Boyd placed 
ii. (big ecus a nasal mite (S. si urn i) from the starling. 
I he many similarities of the two mites Justified [his 
action Dr Womersley (sic) lias suggested (private 
correspondence) that the type species. £ uiafrults, may 
be a nasal mite and that drinking water may be the 
vehicle of transmission. The author is in complete 
agreement, Speleo%nuthus xft'iutus has a hydrophobic 
cuticle, enabling the mite to float on the surface ol ihe 
water, and also is able to run quire rapidly." 

The second Australian species of spcleorrjuttbtd (or 
speleoe nut nine the Spclcognnrhidue me now 
geneially considered as only o( subl'amilv StfltiJS 
Speleognathinae, by recent authors) mite to be 
discovered was described by Womersley 1 1953), who 
erected the genus Hovdaia for Spetecwwfhi** Ml 
Boyd. 1948, and included in tins new genus H tvwdtw 
Womersley I953| found in "mucus under the tongue 
of a frog Lhnnodynastes rasn/aniensis Gunther van* 
in Adelaide, by Miss Laura Madeline Angel-, M.Sc . id 
the Zoology Department, University of Adelaide, while 
.she was searehme for internal parasites. This SpQ 
is now placed as Lawnncutvs angelae <Womer-U'., 
1953) (see Domrow (1961, p, 379>l 

Mi«»s Angel hai informed me (pers. iflinm., IV7<Y| 
ihat the mile was found stuck in the mucus under Ihc 
mnguc i.i a lr"£, <n whose oropharynx she was 
searching for trcmatocle parasites No nitre bfiftdViUUI 
was observed, This male frog had come fnoni Meadow-. 
Mr lolly Ranges, South A.isrmlin, in QCtflbcl 1952 
(coll. I. M. Angel), the ftOg was recorded as 

L ui\murni>riMs var. platycephalus. Miss Angel has 
commented that she has examined many hundreds ol 
Irogs from i he Adelaide ro lower Murray River regions 
without finding any other specimens or speleoguaihmc 
mites. Mi Michael Tyler, Department of Zoology, 
University of Adelaide, advises (pers. comm., 1976) that 
the vai iety fjiurycejthulus is no longer recognized, and 
the name of this species ol Irog remains ai 
/Jmnodynaves tasnuwiensis Gunther. 1858. 

Womersley also included in his genus Bovduiu I he- 
North American species SpCteOgVffthuS striatus 
Crosslcy> 19 V 

Tn rm 1953 paper Womersley declared that & 
microtis had been collected by myself "in moss and also 
on l he surface of water in horse troughs aT C.liu 
Osmond in 1934 and 1935 . • • 

"As all the specimens were females and from the 
habiial on horse Houghs it was thought that in the early 
stages Ihey may have been parasites in the nasal cavities 
of birds or cattle di inking at the troughs. 

Ihc swabbing ol cattle and the examination ol 
birds, however, failed to show any evidence of this." 

A further mite from the Australian speleognatluiK- 
fauna was described by Womersley in 1954, as Boyduia 
derrick! Wometsley, 1954. This was collected upon 
Ratlus assimilis in Queensland, Australia, and was the 
first recorded speleogr.athiue mite from a rodent. At 
the time Womersley believed Ihat the association was 
'"probably accidental M . Subsequent work by Fain 
(195513), Domrow (1961) and others has shown that 
several species ot speleoguaihiites are parasites of 
rodents. This species is now known as 
Parasptdeogtwihopsis derrick! (Womersley. 1954) (sec 
i -in. 1963). Domrow (1961) has recorded this species 
in Queensland from Rut/us conaUts and Rutins raUus, 
as well as Rutins ussimilis. No further reference to 
Speleognathm >amirulis was made by Womersley in that 
paper (1954). which was his last coniribuiion on ihc 

In 1954 < untemari recorded a hurl her species of 
spcleoruailiiiM- < i m I c. SpelQOgrtdthOpSiS gatlh as a new 
genus and species. Irum the nasal cavities ol the 
domestic fowl, Gatlus ^ollus I , a( Asinda, ui the then 
Belgian Congo. In the most reeenl revisions of the 
nomenclature (Fain 1963; Domrow 1969) this speeds 
remains is Spctcnxnoihops'ts galti Coorvman. 1954. 

fain (1955a) described bricflv, in ail addendum, a 
spelragnatfune mile from Ami Ida. as Speieo?notht*s 
bavls fain obtained liom the nasal cavities til 
"■bnvides" (species ol catlk* were noi named). Fain later 
ll956b) described ihis species in more detail. Fain 
C1956a a b) suired thai rhe maxillary and frontal sinuses 
weic die actual rites for the! mites The canle were.slitl 
reh-rrvd io as "bovidc^'. recently (1983), <n response 
to the preseiu author's request, Dr I am Ikin advised 
thai the mito were found in the masillaiy :n\d frontal 
sinuses of Zehu eafilc, i.e. of Has tndicus L. 



I atei Ifl 1956 Lain (1956c) was able to stale that & 
bovis was a synonym of S. australis, from an 
examination of authentic material which I collected 
from Gicn Osmond. Moreover, the previous separation 
was based on inaccuracies in Womerslcy's descriptions 
and figures (Womcrsley's own admission, quoted by 
Fain (1956b, p. 662)), Fain was also able to include a 
statement from myself correcting the attribution of the 
Australian material to moss v and il is also corrected 
in Southeott's (1957) and Domrow's (1961) papers 
referring to speleognathine rnites. 

Drummond and Medley (1964) recorded that on one 
occasion, in November 1961, near Cache, Oklahoma, 
two out of three North American bison. Bison bison 
athabascae (Rhodes), examined had an infestation of 
Spcleognafhus uusiralts in the nasal cavities, with 
evidence of an abnoimal sinus condition, in the lorm 
o( blackened areas of epithelium, and the sinuses filled 
with brownish fluid, 

Thus the early suggestion that the original 
speleognathine discovered, Spek'ognathus atisirolis, 
was an endoparasite upon domestic cattle was 
substantiated by the observations in central Africa, and 
in North America. 

In subsequent years mites of this subfamily have 
been recorded from a wide variety of birds and 
mammals as well as other vertebrates, in several 
continents. In his review of the Spelcognathinae Fain 
(1963) listed over 40 species, divided among 8 genera 
and further subgenera (omitting iMWtvncurus and 
Ratrucorus, placed by him in the subfamily 
Lawrencarinae of the Eicynetidae). Since that tunc 
further nasicolous ereynetid mites have been described 
from Australia- see Do m row (1965, 1969, 1975)— and 
more general reviews have been published by Fain (1969, 
1970a, b, 1971a, b). Fain and Aitken (1969), Fain and 
Hyland (1970, 1975), O'Connor (1978) and Hyland 
(1979). The nomenclature of these mites at present 
uppc;irs in have reached a degree ot stability. 

h is an Interesting point that had it not been for the 
fortuitous circumstance that I used cattle-troughs in 

mv area as a means of sampling and collecting small 
aithiupods, Speleognathm utistmlis might never h;»\e 
been known to occur in Australia at all. Moreover, had 
there not been, in Womersley's estimation, an affmnv 
or resemblance to the mesostigmatic mite genus 
Spelaeorhynchtis Neumann, 1902, it is safe to say that 
the history of the nomenclature of this subfamily of 
miles would have been vastly different. 

The literature of these mites is now extensive, but 
it is not proposed to attempt any general survey here. 
Instead, the purpose ol this paper is to place on record 
a number of early observations upon the times of 
occurrence, and certain details of the behaviour Of the 
mite Speleognathus uusuvits Wornctsley, which have 
either not been recorded at all, or else recorded scantily, 
and in a somewhat corrupted form in the literature. 
It is considered that this is justified, since all the 
Australian observations upon this species have been 
either in the notebooks or memory of one person, the 
present author, for 40 years or more. The mites have 
not been seen in Australia since I939 t and (he 
opportunity of making observations on the species in 
Africa may well be limited for some time to corne. 


(a) Locality 

Over the years 1934 to 1950 (as well as somewhat 
earlier and later), the area studied at Glen Osmond for 
Speleognathus australis consisted of some open 
paddocks upon the lower and north-west slopes o) 
Mount Osmond (Fig. 1) at MR656807 to 655806, at 
a height of 183-191 m (600-625 ft) above mean sea level, 
on map Adelaide, 1939, No. 810, Zone 6, Sheet 
I54M/1V SF & SW (Military Survey of Australia, 
1:63,360). This area is shown in Fig. 2. The two main 
paddocks were separated by a stranded wire fence and 
a roadway (Fig. 3). Both paddocks were used by a Mr 
f.ioldsack for agistment of small numbers of domestic 
cattle (Bos taurus L.) (invariably cows, as no bulls or 

Mt , L (icneral view ol Mourn Osmond, South Australia, I mm Ihe north-west, Iron, a phoiograph on \H August 193ft An ■•! 
Ihe Halter IhiiJ aiound the base Off the mountain has been commonly Milled filen Osmond. The three cattle troughs upon winch 
all specimens ot Speitfofinathus anstr<t/is have been collected in Ausiralia are ai (he middle level ol (he photograph. Trough A is 
tiluatcd somewhat lo the right of the centre, in the dark dump of trees. Troughs Band Care lutihet to the right, behind Ihe tine ot trees 


REC. S. AUST. MUS. 19(13): 201-212 

June. 1986 




[Glea /Ofmiond ^ t °smo^ 

r. House r&W 

te — 



FIG. 2. Map of the sites of the three cattle troughs at Glen Osmond, South Australia, from which all Australian Speteognatnus australis 
Womersley, 1936 have been collected. A, B, C arc the sites of troughs A, B, C respectively. The grid squares arc 1000 yards square, 
or 974 m square. 

steers were included), perhaps 10 to 20 at the most, and 
usually many fewer. Horses were also occasionally kept 
there. Mr Goldsack had no objection to his paddocks 
and cattle-troughs being used for purposes of 
biological surveys, as long as there was no interference 
with or disturbance of stock, and would occasionally 
ask what was being observed upon the cattle-troughs. 
The troughs were of the usual galvanized-iron type 
common in Australia; half-cylindrical, about 45 cm 
across, and with a float-operated cistern to replenish 
the water from the suburban reticulated water supply. 
Of the three troughs studied, one (trough A) (Figs 4 
and 5) was actually on the ground surface, being 
supported against damage from the cattle by being 
banked up with the clayey soil along the sides. Some 
grass and weeds at times grew alongside the troughs, 

but the soil around each of them was largely bare from 
the continual tread of the cattle and horses. The other 
two (troughs B and C) (Figs 6 and 7) were close 
together, to the south-west of trough A, and were 
placed with the bottom of the trough perhaps 15-30 cm 
above the soil. Troughs B and C were supported by 
wooden posts at each end, and trough B also had 
additional supporting posts at about the middle (see 
Figs 6 and 7). Although the soil around troughs B and 
C was as well trodden as that of trough A, some weeds 
managed to grow underneath them and at the ends 
where access to the animals was denied. Each of the 
three tanks had some protective boarding over the 
cistern, to prevent it being damaged by the cattle. This, 
to some extent, limited observation of the water 

FIG. 3. The roadway, fences, and lines of Eucalyptus cfudocalyx separating the paddock of trough A from that of troughs B and C 
Looking southwards from near trough A. Photograph taken in about 1938. 



FIG. 4. Trough A and its immediate surroundings, looking up the valley of the centre of the previous photograph (Fig. 3). One cow 
is present. The trough is on the ground, and is banked up by earth. Photograph taken on 28 January 1938, at about 6 p.m. 

FIG. 5. Trough A, looking to the west; with the same cow at the trough as in Fig. 4. The large trees are Eucalyptus cladocalyx. Photograph 
taken 28 January 1938 at about ft p.m. 

FIG. 6. Trougn B, looking to the north-east. The trough is in an open paddock. Note the straw on the ground, and many patches ot 
dung. The trees are Eucalyptus cladocalyx. Photograph taken 28 January 1938. 

Trough A was overhung by a row of sugar gum trees, 
Eucalyptus cladocalyx, which even in the 1930s must 
have been about 15 m high, and presumably a number 
of the psyllids and other insects found on the surface 
came from the foliage of these trees. The other two 
troughs were also placed about 10 m from a row of 

Eucalyptus cladocalyx bordering the adjacent paddock. 
All of these trees appeared to be part of one planting, 
and according to Gill (1905, p. 5), had been planted 
in 1895. 

At times birds were seen to drink at the troughs, and 
feathers would be found floating in the tanks. There 


REC. S. AUST. MUS. 19(13): 201-212 

June, 1986 

FIG 7. Trough C, near trough B. Photograph taken 12 June 1938. 


























































































— No observation 

.. Mites not recorded, presumably not observed 

* Not all of these mites were taken. When they were numerous I was careful to take only a representative number, to preserve a presumed 
breeding population. 

appears to be little doubt that these troughs were used 
freely by the usual range of birds found in suburban 
Adelaide at the time. The tanks were observed 
occasionally to have large insects such as small 
grasshoppers, floating in the water. It was observed also 
that bees, wasps, and other insects used these troughs 
for drinking, either directly from the surfaces, or from 

No mammals other than cattle and horses were 
observed to drink at the troughs, but dogs and cats 
probably used them occasionally. 

Among the local wild mammal fauna were possums 
(Trichosurus vulpecula), bats, rats, and doubtless 
others; there were no local kangaroos or wallabies. 
Although a few snakes occurred in the vicinity (the 
brown snake, Pseudonaja textilis (Dume'ril and Bibron) 
being occasionally observed), there was no evidence 
over the years that snakes actually came near the 
troughs. However frogs undoubtedly used these waters 
at times. 

(b) Times of Occurrence 

It was my practice for a number of years, to walk 
each week from my home at Glenunga Avenue, 
Glenunga, to the Glen Osmond sites for routine 
collecting. These sites are mentioned in my work (1946) 
on the Erythraeidae, and the location of the three 
cattle-troughs is close to the "second situation" 

mentioned in that paper (p. 7). I recorded my findings 
systematically until 1940, the only exemptions being 
when I was away on holiday. The numbers of mites 
observed over the years 1934 to 1940 are shown in 
Table 1. 

Inspection of Table 1 shows that the numbers of 
mites on the water surfaces of the troughs were 
maximal in autumn to early winter (March to May), 
but there was only one month (October) in which no 
mites were found during the seven-year study. 

Over the years I gained the impression that the most 
likely time to find mites on the surfaces of the troughs 
was during a spell of fine weather, after rain. It should 
be remembered that all the observations were made at 
a time when there was no knowledge of what the host 
animal's identity might be, or in fact if there were a 
host species of animal. 


I propose to quote in this section a number of the 
observations recorded in my notebooks, since similar 
studies have apparently not been made upon this 
species of mite in its only other known localities and 
sites of occurrence, in the nasal cavities of cattle in the 
former Belgian Congo, or of bison on Oklahoma, 
North America. 

28. ii. 1936. There were two animals [Speleognathus aitstra/is] 
found, one on the surface of water on horsetrough A, 

fMSttWKRA AND NATURAL HlMORV Ml SHkt HJGNAtfilS Al \/ A 1 I ' .'.', 


and another on the surface of borset rough C. | Through- 
put niv noies the term horsetrough is often used. 
although CfltifB weir by far the major users of these 
troughs.) The miles seemed 10 he Kjuttfl al home on die 
surf-ice oi i In. water; holh were running quickly over it, 
slopping al limes neat small objects as though in search 
lor inod. Ihc one on tiorseuough C mounted the side 
of the iron trough about 4 em above the level of the wmet 
and then descended. Then it eatue upon solid material 
(floating) it ran over it, but onk for Q few seconds ai 


14, iv 19.16, I revisited Ihe horseiroughs, The miles wert present 

iu abundance on horsctrough B; none was present on 

i I of horsetroughs A and C. On horselrough B 13 

: i were counted; these WCfl all rtl IS dUiCkly WS 

the *aiei sutface, appatfeoUV eithci m seaieh of food 
and/or males, or suitable places for egg-laying. They 
were all i uniting i ai r 1 v quickly, with no noticeable 
dil Terence in their average speed All, io the naked eye, 
wrie of the same sue One mite was observed to leave 
the surface ol Ihe water, then Io mount Ihe sides o! the 
trough to 8 height of aboul 3 cm above the water, then 
Id run along the (rough for about 20 em keeping nearly 
at the same level, then to descend and ro run ever the 
water surface once mote. J again noticed the preference 
of 'the animals tor the waier surface; most ol them did 
not leave the surlace at all except to tun over some 
iloaiing grass etc. This was about 3 o clock in the 
afternoon; there were about 10 cattle neai die trough. 
There were no horses, although last Saturday I here were 
about 4 horses there as well as cattle. 
The last iwo or three days have been line and wairn without 
exception; last Saturday (M,iv IS) was rather POGJ, and 
only one milt: was louud (horsetrough B). This seemed 
to be in a state of torpor oi dead, it was lying on its 
hack 011 the water 
It appears that a warm spell is necessary io bring the |jfjW* 
out; I have noticed this several times. 

Possible ccifdudotis 

1. Miles were all of the same se\ 

2. Miles were seeking to Lay eggs. 

3. i-ront the fact that it has only been found on the water 
surlace ol these three troughs it may be that it is 
parasitic m some stage in ealtlt\ e.g. bronchial 

The continued search for clues as to the life history 

ol Spi'leognaihus ausiruhs led to a number or 

observations on the minutiae of behaviour. As none 

of these has been previously recorded* even though the 

mile has now been described for over 40 years, and as 

i hey have not been do pi icy ted in the scanty 

observations on this species in Africa or North 

America, it seems worthwhile to publish them. 

6. i. 1937, One of the three mites taken on 6a IW was kept 

in D idass jut on |ihc| surface of water foi several days 

V.nious insects were thrown on to the surface but the 

mire was not seen to feed on any Picee|s] of trass etc 

were placed on the water, and although the mite fan 

■ caddy over them it always came to rest on the water 

It can move its legs about 10*15 limes per see. it would 

run near Ihe side of Ihe jar and vibrate its legs rapidly 

wilhoui '.hanging its position. The purpose is tu rne, 

nhsuire, \of the animal could easily move u<? the curved 

sudaee o( the watei to die edge. 

One other til die thru, was drowned hv muucision lor 
abom \ hour. 
13 [J983 Two were found on iroUgh A. These were QUift 
i ' < . wricpur ma rube (without water) 

[and taken on a journey], the day was hut. At night thev 
were found to be dead. 

27JU93?! ihe speed ol Ihe miles on I he mm face ol the water 
in the troughs was estimated •*' 1-3 cm /sec 

2.iii.l937. I hree were seen on trough A. l'hese attacked a fly 
\Muscu) in ihc water of the trough simultaneously [far] 
Ffrod? I To lay| Bffl$3 |lt now seems more likely that the 
fly might have had comae! will. Ofti nasal mucus hum 
;he calllcv and thus provided a suitable aiiraclani for 
the mites.] 

2n ni is+*7 ( Two specimens [were] captured running over the 
surlace ot trough A. These were placed in a tube, on 
Ihe surface of some water from llhe] trough, togeihti 
with aortic of the insects [that had] fallen into the trough, 
and d specimen o( Cvpri\ (Crustacea: Ostracodai from 
the water. After two days the miles were recorded a.-. 
dOl liiant, |audj exhibited no signs o\' life on touching 
with a needle. [They] remained like this for some time. 
The mites remained thus and were almost completely 
decomposed by 20.V.1937. On l4.Viti.IS37 the residual 
scraps 01 nulc skins were removed and shde-mouuled. 

16.iv.1937. One decomposed mite was found on the trough. 

I0.v)(.l937. Another dead mite was found: t'irsi m'ne found 
since l6iv,19J7, 

Jf>,\ii.I937. One mite lound on trough A, tipped OW on its 
back, ll moved iLs legs actively when first found* bul 
was feeble a little laiet. Mite taken. 

17jtii.l937. Another mile found and taken, aciive- tidied he 

27.xh.iy37. Trough A was examined al 4-5 p.m. Three 
specimens were present One was dead and decomposed, 
the other two were active, plump, running actively over 
the surface ol rhe waier. the two came together, circled 
rapidly, and met (anterior end to antenoi end). They 
were like this for a second or two, moving their legs all 
iIk- lime, then they separated by 2 or 3 mm, and were 
blown apart by the wind . . 

The trough Was re-evamnted at 8 pan., when the 
decomposed mite was as before One was moving its legs 
slowly; one was dormant with its legs drawn in under it 
On trough C at 4-5 p. in there was one very plump 
and active mite at fhe surface o\ the waier. At 8 p^p. 
ih^ (presumably same) mite was floating, tipped over 
on its back. 
On l.i.1938 I again examined the three (roughs al 

8 p.m.. looking for evidence of possible nocturnal 

characteristics oi the mite I made the following note: 

I he evidence of the past lew days suggests strongly 

thai the Spet. does not leave the water by climbing « ' 

|of] the trough— it is possible that it leaves it by means 

of the cattle drinking there. 

On 2.L1938 the troughs were examined at about 7pm 
(sunset). On trough A there was one Spelcognathus, \ cry- 
plump, and running quickly over the water. This I took 
Troughs B and C had none. The animal was observed 
on water in a dish. It moves by moving only the la-i *^ 
joints [i.e. segments] of its legs. The tarsus is practicallv 
at a right angle to the water surface when at rest, or when 
moving It docs not move by moving its trochanter on 
the coxa, but by moving lis tarsus and metatarsus [i.e. 
tibia] backwards and forwards in a nior? or less vertical 
pf&W. The animal was drawn in outline, and in its main 
character is- ics. and then killed with Carnoy^s lluid, No 
trace ol eggs could be seen in the body. Shape is as 
figured, widest anteriorly, narrower posteriorly, with 
sides becoming approximately parallel. (Tiguic is 
shown in Fig. 8). 

Spelcognaihu.N is delicate and cannot withstand much 
handling or shaking about— either on water or dry. 


REC. S. AUST. MUS. 19(13): 201-212 

June, 1986 











h 'd,Or^j( 

-pefep ( 

n6 k. ^*«*i 
j A US --o' " ?// i : v$^. 

ii ■ ' 

■The tffth'M'h'? p4 *£ «^i!c %.Kk.jL* 

FIG. 8. Speteognathus australis Womersley. Living specimen from Glen Osmond, South Australia, 2 January 1938. Drawn freehand 
through the microscope, on the surface of water. The main lines have been darkened with pencil for reproduction purposes, otherwise 
the drawing is untouched. The reticular patterning of the legs is omitted. 

On 28.L1938 I photographed troughs A and B (see 

Figs 4-7) and recorded in my notebooks on that date: 

The cattle are shifted about constantly in the paddocks 

containing troughs A, B and C, but rarely are any of 

the paddocks containing the troughs without cattle for 

more than a few days. 

A few further observations were made during 1939 

and 1940, but I find from my notebooks that all records 

for 1940 were negative, and the last living mites seen 

by me were collected in December 1939. (Previous 

statements implying that the mites were seen as late as 

1941 were based on faulty recollection.) 

On 29.U939 I observed four mites on trough A, but 
none on B or C. One of the four was alive; three were 
dead (one apparently not long dead, one with legs 
flexed under it, and one decomposed). I made the 
following comments in my notebooks: 

Conclusions: The hot spell has brought out 
Speteognathus , following as it does fairly wet 
weather . . . Note: Dead Speleognathi are found on the 

1 observed the living Speteognathus for about half an 
hour [on the trough]. When the . . . Gust dead) 
Speteognathus was put in its way it ran over it without 



stopping The animal seems to give no preference to light 
or shade. Kan for a length of a metre (with frequent 
stoppings, arid many turns, and doublings back, and 
assisted by the wind, and water Currents. Slops varied 
I'rom 0-10 seconds, rarely the latter, usually about 3 

It was put near the edge of the trough, but did not 
go up ar all. Also, it took no notice of algae just below 
the surface of the water. 

Wasps, bees [are) observed drinking here. Cows drink 
here, and dogs sometimes; and birds (magpies) [This 
means the whire-backed magpie of south-eastern 
Australia, Gynworhina hypoleuca Gould, These 
frequently nest in tall trees, such as the rows of planted 
Eucalyptus clariocalyx bordering the paddocks 
containing the troughs. | live here), and some bird 
feathers [were] seen in |the| trough. Many ants [are] 
about; some fall in the water. One butterfly (Danaus 
archippus) [Now Durtaus ptexippus plexippus (I.. 1758) 
(sec Common and Waterhouse, I972,p, 221)] [was] seen 

Troughs B and C were swarming with Cypris. Troughs 

A, B and C were swarming wuh nemaiidiform (almost) 

larvae. Small bugs run over (he surface of the water. 

Water dirty, much alga present. 

A sketch was made of the path of the Spe/eognuthns 

on the water, over a small area, to indicate its many 

twists and turns. This is reproduced in Fig. 9. 


HCJ. 9. Drawing made on 19 January 1039, oj the path of a 
specimen o\ Spe/eoyjmihus justralts tim a limited area of the 
w;iit-r surface t>t trough A at Glen Osmond, South Australia, 
reproduced at Original ttzc Tne wind direction, and water 
current dirEttiOnS fcJ* S&OW0 It is apparent Irom thil sketch 
that the mite was not under the control of either the wind e>r 
water cm rents, and could move al will upon the water. 

On 9.i\.1939 I observed 25 of these mites upon the 
surface of trough B; there were none on troughs A and 

All of these were dortnaru with the legs urongty 
fii'xcd Not one was seen to be in a state of 
decomposition. Not one was seen to give any movement 
a| all, although several were stimulated by touching and 

several were taken (7 in all) — 5 were laken as | hey were, 

2 others put in alcohol — not seen to move M all). The 

mites were in any position e.g. tilted on their posterior 

ends, their sides, or in normal position. None was on 

its back. The occurrence is most remarkable . . . the only 

sunny day recently is today ... no cattle were drinking 

at the troughs, or present in the paddocks ... on 

SL1VJ939, The five unpreserved mites were examined 6 

hours later, but were unchanged, and were then preserved 

for histological study. 

Observations continued, but did not throw any 

further light on the presumed host animals or ul'e- 

history of these mites. 

By 1940 I was systematically noting the various 
species of birds seen in the vicinity of the troughs, but 
without drawing any significant conclusions. 

During 1941 1 was able to make only a few 
observations, and thereafter, owing to military duties* 
1 was not able to inspect the area again until 1946. By 
then troughs A and B had been removed. Trough C 
remarried and was examined at irregular intervals from 
1946 to 1949, and occasionally later, to I&53, but no 
further Spelvognathus was found. 


The possibility that these miles were endoparasites 
of the nasal or other respiratory passages of mammals 
and birds drinking at the troughs was entertained quite 
early. Since the only residual local marsupials were 
arboreal ones, such as Trichosurus vutpecuia ana 
smaller species, it did not seem likely that marsupials 
would prove to be the hosts. The animals drinking most 
commonly a) the troughs were caitle, with a smaller 
population of horses, but It would seem likely prima 
facte, that if such mites were parasites of the nasal 
spaces of cattle and horses, they could hardly fail to 
have heen detected in Europe, the obvious source of 
the Australian domestic breeds of cattle and horses; 
not to mention the other domestic animals in South 
Australia, such as dogs, cats, and so on. Similar 
considerations applied to introduced rodents. Bats 
appeared to be another possibility, but rather a remote 
one, although insectivorous bats were not uncommon 
in the neighbourhood, even if mainly crepuscular or 

Other possibilities that were considered, cither then 
or later, were thai the mites could be tracheal parasites 
of water insccis occasionally seen upon (he ttoughs. 
Conceivably also small gastropods might serve as hosts, 
in the same way as Rkcardoella utilizes the common 
introduced slugs in South Australia, because of the 
obvious resemblance between Speleognuthus and 
Riccardoetta. However, no solid evidence was found to 
support seriously any suggestions of a non-vertebrate 

Since the mites were reasonably common at times, 
and since the cattle were the commonest large mammal 
niili/ing the troughs, it seemed worthwhile to 
investigate further the possible role of the cattle 


REC, S. AUSr. MUS. I»<U>: 201-212 

J«*e, /9fiffi 

My noiebooks record; 
20.V.1936. Cattle drinking ad (roughs A. Band C ai Cilcn 
Osmond. South Australia, had their nostrils swabbed 
No Speieog. were obtained. 

This is not surprising, and probably tht result does 

501 mean very much, since more ihan a month had 

passed since any of these mites had been seen on the 

troughs, ftfestl performed by C.S.I.R, officers, at 

suggestion of H. Womersley (To them, that isM 

I was not a witness to the event, the information 

having come from Womersley (1953, p. 82) who referred 

lo it briefly. 

After i his negative result, no further attempt was 
made to locate the mites in the nostrils of the cattle. 
Subsequently Lhe possible relationships remained 
matters of speculation, as the various new species'of 
speleognathines were discovered in North America, 
Africa, and other continents. The next species to be 
discovered was by Elizabeth Boyd (1948), in North 
American birds. 

At the end of 1938, under the existing means of 
study, I had summarized the possibilities of host- 
relationships in my notebooks as follows: 

(a) Nature of the appearances of the animal on the water 
sun . 

1. This cannot be foiruilous — from ease of its 

2, There are no streams left la this locality — even hj 
winter they do not run (although they do contain 
water for a while)— and only remain with water 
in them for a short time. The troughs are from 
10-25 m from lhe creek bed 

(b) Fauna which might be related 

(1) Non-aquatic 

Birds (the word especially added in ) 
pencil at some later date, and the i 
word birds underlined) ) drinl 

Cattle ) J,e, e 

Insects including wasps, bees, bugs, etc 

(2) Aquatic ) 
Trough contains a rich fauna of Cypris, In Dec 1938 
many nematodes (?) were present. (There is a good 
collection of filamentous algae in each trough) 

(c) Flora which might be related 

(1) Non-aquatic— eucaJypts, grass, weeds 

(2) Aquatic— green algae mainly ISptfogy/a 
Addendum 111939 

Only 3 Speleog. (werel seen m 1938 {and caught), (cp, 
J935 with 26, 1936 with 23, 1937 with 34 (approx. figs.). 

It may be that the continual taking of the Spel. has 
reduced their number considerably 

That surmise was contradicted by the finding of a 
large number in 1939; in fact the count of 25 for March 
1939 was the highest for any individual month over the 
whole period of the observation. 


Over the last 40 years I have looked systematically 
for Speteognathus austratis (and related mites') when 
travelling in Australia and Papua-New Guinea No 
cattle-trough encountered which contained water was 
ever left uninspected. Despite searches in Papua- New 
Guinea (two sojourns) and in every Australian state 

except Western Australia,, none has been found. 
Additionally, a search has been made for intranasal 
mites in birds and other vertebrates from 1942 onwards, 
as opportunity permitted, but again all examinations 
failed to reveal any speleognaihinc mites. 

After lhe discovery of S. sturni by Elizabeth Boyd 
(194S) in North America, 1 decided to make more 
.systematic and frequent efforts to find these mites at 
bird drinking sites. I erected two troughs at my then 
home at Unley Park, a southern suburb of Adelaide, 
by cutting a 44-galIon drum in halves longitudinally. 
One. (D) was erected about a metre above ground level, 
on the stump of a cypress tree, and the other (E) was 
erected on a wooden frame about two-thirds of a metre 
above ground level amidst Irecs, bushes and other 
vegetation. Birds were encouraged by suitable feeding 
to drink at the troughs. Trough D was observed 
regularly for nearly 4 years, and trough L for over 3 
years. Observations were made daily, at times oftener. 
over much of this period. Despite these efforts, 
however, no speleognathines were found 

During 1952 and 1953 1 carried out systematic 
observaiions of the horse-troughs still located around 
the streets, parks and other open spaces around 
Adelaide. Thus 40 observations were made in March- 
Juno, I953 f and observations continued into 1954, 
when lhey were abandoned. Again, no speleognathines 
were found. 


It may reasonably be concluded from these many 
hundreds of observations that Speleognathus austratis 
must be an uncommon species in Australia, If it were 
a common parasite of cattle, it is reasonable to believe 
lhat it would occasionally come to the notice of 
veterinarians. There is, of course, no evidence at present 
that this species of mite is concerned with disease 
transmission in any way, ot in fact that it causes serious 
harm to its hosts, Nevertheless, the lesions recorded 
briefly in one specimen of North American bison by 
Drumrnond and Medley (1964) show that this species 
of mite is capable of causing some tissue damage. 
Presumably the mechanism is similar to Unit in, for 
example, Rkxardoelta timueum (Schrank) (Ereynetidae) 
which feeds on slugs and is capable of killing them 
under laboratory conditions of heavy infestations 
(Baker 1970a, b). Baker (I97J) studied the ereynelid 
mite Xenoparcarus africanus Fain, Baker and T'msley, 
1969, which lives in the nasal passages of the African 
clawed toad Xenopus laevis Daudin, and demonstrated 
blood in the gut by histochemical tests. Both of these 
species of mites have mouthparts which appear capable 
of penetrating epithelium. 

Baker (1973) discussed this further and commented 
fp. 51); 'The feeding of ereynelid mites, involving the 
production of a stylostome. appears to be basically 
similar to that already described in such trombiculid 
miles ?js Neotrombieula zachvatkini Schlugcr and 
Ttambtcula antumnatis Shaw . . . ". 



Other groups oi' invertebrates have exploited the 
same niches, such as various mesostigrnatid mites, and 
the family Trombiculidae (sensit lato) of the 

The second major question which a student Of tin? 
distribution of Speleognathus australis will ask, is how 
can we explain Ihe grcally separated distribution of the 
species, known only from three recorded localities: 
Glen Osmond in South Australia, Asfrida in the former 
Belgian Congo, and Oklahoma in North America. In 
the African and North American cases there was 
evidence ol nasal parasitization of a bovid, while in 
the Australian case there was evidence pointing to an 
association with cattle. Only one species is now 
classified in the genus Speleognathus; all Other 
speleognathines, from a wide variety of vertebrate 
hQSfS, are now placed in other genera (Fain 1963). 

We may reasonably accept that Speteogncrfhus 
australis is a bovid-adaptcd nasicolous species. 

The problem remains* however, of its extremely 
disjunct distribution as at present known. Although 
I stated (1963, p. 328) that this was ^easily explicable 
since a number of the original cattle of Australia were 
Of African origin", it now seems that thai explanation 
was too facile, particularly in view of the finding of 
Speleognathus australis in North America, In any case 
the proposal did not explain the lack of reports of the 
rtiftfi in southern Africa, nor from New South Wales 
and other parts of Australia. 

At this point we have to leave the study of its biology 
and distribution with much of the mite's natural history 
in Australia at last accurately recorded m some detail 
but with some major ecological questions snll 


BAKER, K ft, 1970a Studies on the lile history of Hierardoe/ta 
iiinacurtt (Schrujik) (Acari-Trombidil'ornieM ./. /tat. Hist 4: 


HAkl-R. R. A, 1970b. JTtt iYmkI or Rtccunh^Hu iimacum (Sehrank) 
-AeaiiTtombidiiowic*. and its rclaOonship svnh pulmonale 
molluscs J. nut- tli\t, 4 $21*530. 

bam K, K. A. 197|. Observations on aspects ol nuiriiion in 

Xnn>[nj< urns afrnunns (KrcviicOdae* TYonihiditormes). J. ntcti. 

Entomah 8: 307 313. 
UAKFR, R. A. 197.*. Ndlei en ihe inltinal anatomy, the food 

requirements and development in die family Dcyneiidac 

( lionibidifoimcs). -hart >U)}; i a IS: 45 S5. 
BOYD. ) li/abcih, M W4K. A new IHilfi bom ihe H»prWl©r> nact 

ol* ihe slarling (Aearhia. SpcIcognaOitdac). Pr<n Eni SQi 

Washington 50: 9 14. 
COMMON, 1 F. a, iind WATLRHOUSF. I- W2. "BuiUrrHies 

0{ Australia" Anyut and Robcrhon, Sydnev. 
(.'OONHM\N, |. 1954. Sui uti acaiicn nouveau, parasitedes fosses 

nasales des oiscaux el rcmarque* »*nr I* boniUe des 

Speleoynaltndae. Ann Para.sitoI. hum. comp. 29; 426-43?. 
CROSSLL V IX A . It, IT--. Fwo new n;r,:d nuics Ir.irn columbttorm 

hirds. / Puntstlol 38: 385-390. 
|>OMROW. R- 1961 The laimlv Speleognathidae in Australia 

(Awarinaj PrCa 1 torn. 8ti< v.s u 9$ .vm-iki 

DOM ROW, R- 1965. Three spelccgnaihid miles from Ausir.-duin 

hirds, \varninfiu 7* 43-48. 
DOMROW, R. |H6V. The nasal FlfltejOJ QUteiftlftlKJ birds (Atari: 

Derma nvssidae, Erevnebdae, and tpidcrmoplidac). Proc, Linn. 

\,,r >\SM 9Y 297-426. 
DOMROW, R. 1975. A new species of Par ( t\petco^nuthi>p\ts Fain 

lAviirr CrcvMct.iLLu.-j hum an Australian dasyurid marsupial. I 
tttl lot SOU. 14: 97-99. 
I iKi MM* tM> R. O., and MEPl BY, J. C. 196*. Occunw.c ol 

tyeJcojwi!fhusaustnillBVfofi\9Tt\ty (Acarftia: 8peieagnatnfcJAc) 

iti Ihe nasal pottages ol bison. 7, Pufaslfoi Sty ^5. 
»AIN, A. lysSn suf un nouvel aearien para&iu des'. nasalts 

de In peidriA nu Rtiaiiu-Uiundi: lioydata pternistis n. Jp. 

(SpClcotinMlllnhic). KVt. ZQOl Rttt- l/Jk S2: 144-140, 

TAIN, A. 1955b. Sur le paratfil&nie dqs Tosses nasales lIkv tcs 
mummireres el les oiseaux pai Icn aeariens clc l« famill** 
Spflrtn'.'tiJihidae (Acji ma). Oescripttort d'une esp^ee nouvellc 
chcz k'i ehaiive-sottr^. Ann Sac, Bet$ Mi'd. Trop. 35: 659-700. 

IMN, A. I956it, U-s Mraruns iIl* la laimllt: Sftck-DKiujlhidae 
Womcrslcy an Ruanda TJrandi (CoHgU T*elge>. Rr\ /ittif. Rot. 

fl/r. S5: L7-«. 
IAIN A. 1956b Presence d'aeanensde la famillc Spelcetgnaiht<lac 
Womersley dam les losses nasalcs dt- mammiferes. 
lie iit.i'i espicei nouveiics Ann Purusuot. hum. comfy 30 

FAIN. A. (956c. Nouvelles obsorvaiions ^ur les ae-ariens de to laimll.- 

Speleognaihidac paiasiies dtS Tosses nasales ehd le* hain-.r n- 

les oiseaux el les maniiiiifcres Ann. Para^itvi hi/rn <i>mf>, .H 

643 -662. 
TAIN. A 1957. Siir la petftKM sys|rniali(pie dr h'krerdvella cweri 

Lawrenec 1952 et de Boydata angetae Womcrsley 1953. 

Reinamenteni de la famillc hreyiiL-tidae Ondenuns 1931 Hov. 

AxtL Hot. a/n 53. ;M9-252. 
hAlfv, A. |9fi3. ChaetoiaMe et classification Cies Spcleoenathinao 

(Aeanna. rror.ihidd(tnnes) Bull Inst. faqy. Set Ntll Bdv W: 

FAIN, \. (96^. Nouvelles Spelcognath'mae paiasiies nasieolcs 

d'oiseaux (Aearinj: lrombidiformes). Rev. /.ool. hot. afr 80: 

TAIN, A 1970a Nomenclature Ues poils idiosomaux el de.scnpiion 

de rrois C$pCCe$ moii^cIIcs dans la farm lie Ltevneiidac 

(TrOmbidifnrnieM. Acamfo^ta 12: 313-325. 
FAIN, A (V70K Aeariens navicolcs tf'otecaui el dc mammiTerrs tie 

Br&il IV Noitveaux 1 reynetidae (lri>mbiditoruu'.) el 

I'lM-hinoplidaL |Sar..opl.loi <ncs> de la region de Helorii (Nurd 

Bfesil). Acvroloyju 12 326 -1*8- 
1-AIN. A- 197U- Nolcs sur les spclcognHlhines parasites nasicolcs 

dc mamnulercs (tre>iietidfle: Tmmbidiformes). Acawtogia IZ: 

509-5 >|. 
FAIN. A. 1971b. (/led hstc dos esptves dujaenre Hovdatu Womerslcy 

(Ltevnciidae: TVombidifornics). Acamiogw U: 98 112. 
TAIN. A., and AITKbN, i. Fl- G i^®, Aeariens nasieoles 

el des mammiferes flu Brcsil. It. FireYTietidat dc la region de 

Belcm (Nord Hre'sil). Bull. Ann, Sor fifty. Fntarn. hetg. 1«S: 


FAIN, A., and HY LAND. K. [• 1970. Aeariens nasieoles des Qiw*U< 

du Mcxiquc- 111. hundks l-.reynctidot* ct llirbinoplidae. Bull. 

Ann. Sot: Hoy. Pntom. gtfg. 106: 37-46. 
FAIN. A,, and HYI ANO, k. E 1975. Spcleognathinae colleeicd Oom 

birds in North America (Aeaiina: Lreyneiidae). J. Aen' York 

Eni. Soc. 83; :<).V20K 
GILI , t, IMS, 'The HtMoiV -ind Tupu^rapl.v gl GlfiH CKn.iuid" 

Vnidon and Pritchard, Adelaide 
HY1.AND, K. C. 1974. Speeilu'Hv nnd pa.allcl hosl-para.siieesv»luiiutl 

in ihe Turbinopiidac. Cylodiiidac arid FacMiciidae living miht 

respiratory passages ol birds In "Keccnt Advances in 

Acamkiyv". Vol II (J. G. Rodriguez, ed.). pp. tt»3<369 Anid.-niu 

Press. New York 
I AVVKENCE, F<. F. 1952 A new paraiitie mi(e Itom ihe nasal chviMl-s 

ol the South African toad. Buj'o regularts Reuss. Prm , /ool 

Sni London 111; 747 732. 
OVONNOR, B. Mi I97S. A redennpiion erf ^peieo/ofivns 

>nirhi*rn\i\ (FordJ IV. ._ornb. (Aean Fivyuetidaul, a nusa\ tnitC 


REC. S. AUST. MUS. 19(13): 201-212 

June, 1986 

of microtine rodents, with comments on generic relationships 
in the Speleognathinae. J. New York Ent. Soc. 86: 123-129. 

SOUTHCOTT, R. V. 1946. Studies on Australian Erythraeidae 
(Acarina). Proc. Linn. Soc. N.S.W. 71: 6-48. 

SOUTHCOTT, R. V. 1957. Description of a new Australian 
raphignathoid mite, with remarks on the classification of the 
Trombidiformes (Acarina). Proc. Linn. Soc. N.S.W. 81: 306-312. 

SOUTHCOTT, R. V. 1963. Obituary notice, Herbert Womersley 
(1889-1962). Acarologia 5: 323-334. 

WOMERSLEY, H. 1936. On a new family of Acarina, with 

description of a new genus and species. Ann. Mag. nat. Hist. 

(10) 18: 312-315. 
WOMERSLEY, H. 1953. A new genus and species of 

Speleognathidae (Acarina) from South Australia. Trans. R. Soc. 

S. Aust. 76: 82-84. 
WOMERSLEY, H. 1954. Another new species of Boydaia 

(Speleognathidae: Acarina) from Australia. Trans. R. Soc. S. 

Aust. 77: 65-66. 



Odontacarus veitchi sp. nov. (Acarina: Trombiculidae) larva from central Queensland is described. 
The classificatory key to the larval Odontacarus of the Australian, New Guinean and south-east 
Asian region is revised. 




Honorary Research Associate* South Australian Museum, North Terrace, Adelaide, South Australia 5000 

(Manuscript accepted 13 January 1986) 


SOUTHCOTT, R. V. 1986. Description of Odomacarus veitchi sp. 
nov. (Acarina: Trombiculidae). Red S. Ausr. Mas, 19(14): 

Odontacarus veitchi sp. nov. (Acarina; 
Trombiculidae) larva from central Queensland is 
described. The classificatory key to the larval 
Odontacarus of the Australian, New Guincan and 
south-east Asian region is revised, 


The author recently (1986) described various stadia 
in The life history of Odontacarus swani Southcott, 
1986, and revised the classification of the larvae of 
Odontacarus Ewing. 1929, for Australia and adjacent 
parts of the southwest Pacific area. Difficulties in the 
placement of O, athertonensis (Womersley, 1945) and 
related species were referred to. In a cognate* study 
detailed statistical analyses of morphological features 
of this group of larvae were presented (Veitch and 
Southcott, 1984), which showed significant metric 
differences between the species presenting classification 
difficulties: Q mccullochi (Womersley, 1944), O, 
athertonensis (Worn.), O. "species S"T (now O. swani 
Southcott), as well as some further specimens. 

This last-named group, from Queensland and 
Papua-New Guinea, included a few larvae in poor 
condition and hence unsuitable for detailed 
description, but also a series of six specimens from 
central Queensland (identification numbers ACB733A- 
F) of which five specimens were in good condition, 
suitable for detailed measurements and description. 
These had been considered as conspecific with O. 
athertonensis larvae by Womersley (slide 
identifications), and by Goff (1979), Re-examination 
of this series shows that it represents a separate species, 
which can be distinguished from other species of 
Odontacarus (subgenus Leogonius Vercammen- 
Grandjean, 1968), particularly by leg metric characters. 

All measurements are in /-im, unless stated otherwise 
Seta and other terminology is as given by Southcott 
(1986) and Veitch and Southcott (1984), 

* n was originally planned that that study would follow the paper 
here nhown as Southcott (1986). However, unforeseen publication 
schedules reversed the order of appearances. 

T Unrelated to an undeseribed species mentioned as V by 
Vercammen-Orandjean (l%8, p 121). 

Odontacarus veitchi sp. nov. 
(Figs IA-3, 2A-B) 

Description of Holotype larva, N 198 1 390 

Colour in life not recorded, presumably red. Length 
of idiosoma (mounted on slide) 237, width 203; total 
length of animal from tip of cheliceral fangs to 
posterior pole of idiosoma 328. 

Dorsal scutum slightly wider than long (nasus 
included); nasus well- developed, narrow, with rounded 
tip and slightly sinuous sides, meeting the body of 
shield at right angles. Anterolateral angles of scutum 
rounded, lateral borders slightly concave, running lo 
rounded posterolateral angles, posterolateral borders 
concave, posterior pole evenly rounded, projecting, AM 
scutalae tapering, slightly pointed, lightly setulose; AL 
and PL scutalae similar. Sensillary setae normal, well 
setulose in distal half Sensillary sockets slightly 
anterior to level of bases of PL setae, set slightly 

Standard and other metric data for type series as in 
Tibfc i 

Eyes 2 »2, sessile, oval, conjoined, near PL angles 
of scutum. Maximum diameter of anterior eye 18, 
posterior 13, 

Dorsal idiosomalae normal, slightly tapering, lightly 
setulose, pointed at tip, arranged 2 (*post-humeral$') 1 
6, then in vague rows across dorsum, of ca 10, total 58. 

Ventral surface of idiosoma with a pair of pointed, 
setulose setae between coxae III, ca 36 long. Venter of 
opisthosoma bears 48 further setae, 24-36 long, with 
outstanding setules; posterior setae longer and slightly 
blunted, similar to posterior dorsal idiosomalae. Anus 
(uroporus) ova], 23 long by 15 wide; about 23 setae 
anterior lo mid-level of uroporus, and 25 setae 
posterior to same level. Urstigma oval, in normal site 
between the contiguous coxa I and II of each side, 18 
long by 14 wide, 

Coxalae 2, 1, 1. AU coxalae long, well setulose, 
tapering to a fine point. Lateral coxala 1 60 long, medial 
coxala I 60, coxala II 51, coxala III ca 55. 

Legs well-developed, normal, 1 416 long, I! 368, 
111 430 (all lengths include coxae and claws). Leg 
scobalae normal, pointed, well setulose. Scobalar 
formulae (including mastalae): trochanters I, I, 1, 
femora 6, 5, 4, genua 4, 4, 5, tibiae 7, 6, 6, tarsi 20, 
17, 13. 


R£C. S. AUST. MUS. 19(14): 213-217 

June, 1986 





\ \ 






/ I 





FIG. 1. Odontacarus veitchi sp. nov. A-E Larva, holotype, N 1981 390. A Dorsal view, legs on left omitted beyond trochanters. B Tips 
of chclieerae, seen in transparency. C, D Dorsal idiosomal setae (c, d in A, respectively). E Ventral idiosomal seta (e in Fig. 2a). All 
figures to nearest scale. F Diagram to show metric characters of dorsal shield. The symbol A indicates that this seta is shown in both 
the dorsal and ventral figures for this mite. 

Leg specialized setae as follows: SoGeI.32d (24 long), 
VsGel.72d (4), SoGeI.72pd (27), SoTiI.55d (18), 
VflU82d (4), S6TiI.85d (16) i.e. slightly distal, also 
anterior, to VsTil. 

SoGeII.25d (20), VsGell.64d (3), SoTiII.36d (16), 
SoTiII.88d (15), SoGeII1.27d (22), SoTill!.35d (20). 

Both tarsus I and II bear a large central dorsal 
solenoidala, I 16 long, II 18 long. On tarsus I is famala 
FaTal, 3 long, level with proximal SoTal. On tarsus II 

is FaTall, 4 long, proximal to SoTall (see Fig. 1A). Tibia 
III has two long mastalae (mastisetae); tarsus III has 
one mastala. Pretarsal formula 1,1,0. 

Tarsus I 91 long by 30 across, II 74x27, III 97x24 
(tarsal lengths exclude claws and pedicle). Tarsal claws 
normal, falciform, slender, with strong setulcs 
(onychotrichs) on anterior and posterior; middle claw 
longer and more slender than neolaterals, with weak 



100 J 

FJG, 2. Odontacarus veitchi sp. nov., larva. A Veniral aspect of hoiotype; legs on left omitted beyond trochanters. B Ventral view of 
palp of paratype NI981385. (Both to nearest scale). 

Gnathosoma normal, well developed. Combined 
chelicerae bases ca 80 across; chelicerae 91 long from 
tip of fangs to posterior pole of bases. Cheliceral fangs 
stout, curved, pointed, with 3 or 4 strong retrorse teeth 
along concave, flexor ( = dorsal) edge, and 2-5 blunted 
denticles along convex, extensor ( = ventral) edge. 
Galeala 30 long, with one faint setule. Gnathobasal 
setae curved, pointed, 25 long, with several long setules. 

Palpi normal, strong. Palpal setal formula 1, 1, 3, 
8, with palpal setal formula B, B, BNN, So + 7B. Palpal 

tibial claw trifurcate. No supracoxalae to legs or 

Material examined 

Queensland: Mt. Jukes, 6.ix.l951, 6 larvae, collected 
on card, E. H. Derrick. Hoiotype with South 
Australian Museum registration number N1981390; it 
bears on the right hand label "Acomatacarus 
athertonensis Worn." and the locality information given 
above, also "G197". The word "Paratype", which was 


REC. S. AUST. MUS. 19(14): 213-217 

June, 1986 

























































































































Sens 51 






























































Gel 11 















1. 00 







l.i \ 






* See the criteria given in footnote to Table 6 in Southcott (1986). 

f Another estimate of this variate is 82 ^m (see preceding footnote, and Veitch and Southcott (1984). 

clearly in error, has been obliterated with white ink. 
On the left hand label is written "Was marked 
l PARATYPE'/ACB133F/Odontacarus veitchi 
Southcott HOLOTYPE/N1981390.". 

The five paratypes, from the same source and 
collector, have similar labels, with serial numbers on 
them as follows: N1981386, ACB733B, G193; N1981387, 
ACB733C, G197; N1981388, ACB733D, G196; 
N1981389, ACB733E, G206; N1981385, ACB733A, 
G192. The unofficial (personal) registration numbers 
with an ACB prefix have been used by me in previous 
publications (Veitch and Southcott, 1984; Southcott, 

Systematic position 

To distinguish this new species, criterion number 17 
in my key (Southcott, 1986, p. 180) should be altered 
to the following: 

Longer dorsal idiosomal setae 58-66 long. 
Genu I 54-59 long. Tibia III 65-71 long. 

Palpal setal formula B, B, BNN So + 7B 

, . . Q veitchi sp. nov. 


Further details of the analyses of the statistical data 
for this and related species are given in Veitch and 
Southcott (1984). 


The species is dedicated to Mr L. G. Veitch, CSIRO 
Division of Mathematical Statistics, in recognition of 
his assistance in the metric studies of Australian 
trombidioid mites. 

17 Ventral opisthosomal setae 48-54 in number . . 


Ventral opisthosomal setae in range of 35-45 
in number 18 

17A Longer dorsal idiosomal setae 42-56 long. 
Genu I 42-49 long. Tibia 111 57-65 long. 
Palpal setal formula B, B, BBB, So + 7B . . . 


1 thank the South Australian Museum for access to 
these specimens. 

Thanks are due to the National Health and Medical 
Research Council, Commonwealth of Australia, for 

O. athertonensis (Womersley) assistance. 




EWING, H. E. 1929. A manual of external parasites. (Charles C. 

Thomas: Springfield, Illinois and Baltimore, Maryland). 
GOFF, M. L. 1979. The genus Odontacarus (Acari: Trombiculidae) 

in New Guinea, with descriptions of four new species. J. tned. 

EntomoL 15: 143-154. 
SOUTHCOTT, R. V. 1986. The genus Odontacarus (Acarina: 

Trombiculidae). II. Observations on the life history and 

morphology of Odontacarus swani n. sp., and related forms. 

Rec. S. Aust. Mus. t9: 169-200. 

VEITCH, L. G., and SOUTHCOTT, R. V. 1984. Discriminant 
analysis using metric data from larval specimens of three species 
of Odontacarus (Acarina: Trombiculidae). Aust. J. Zool. 32: 

VERCAMMEN-GRANDJEAN, P. H. 1968. The chigger mites of 
the Far East (Acarina: Trombiculidae & L^euwenhoekiidae). An 
illustrated key and a synopsis; some new tribes, genefa and 
subgenera. (U.S. Army Medical Research and Development 
Command: Washing D.C. 20315). 

WOMERSLEY, H. 1944. Notes on and additions to the 
Trombiculinae and Leeuwenhoekiinae (Acarina) of Australia and 
New Guinea. Trans. R. Soc. S. Aust. 68: 82-112. 

WOMERSLEY, H. 1945. Acarina of Australia and New Guinea. The 
family Leeuwenhoekiidae. Trans. R. Soc. S. Aust. 69: 96-113. 




AUGUST 1986 



North Terrace, Adelaide 
South Australia 5000 


byPatriciaM. Mawson, L. Madeline Angel and S. J. Edmonds 


This checklist includes all original published records (to the end of 1983) of the helminths occurring 
in Australian birds, as well as undescribed specimens known to the authors to be held in museums 
or other institutions. 




Honorary Research Associates, South Australian Museum, North Terrace, Adelaide, South Australia 5000 

{Manuscript accepted 7 November, 1985) 


MAWSON, P.M., ANGEL, L.M , timj EDMONDS, S.J. 1^86. A 
checklist of (he helminths from Austialian bitfta. Re*. S. Aust. 
Mus, 1^(15): 219-325. 

This checklist includes all original published records 
(to the end of 1983) of the helminths occurring in 
Australian birds, as well as undesenbed specimens 
known to the authors to be held in museums ar other 

Taxonomic lists given arc (1) bird hosts, numbered 
consecutively, with the helminths from each bird noted 
below it, with ihe locality of collection and the 
authority for each record; and (2) helminths, with the 
bird hosts listed below each helminth species. 

There are also three alphabetical lists, ont of the bird 
families, one of the bird species and one of the 
helminth species, all referring by number to the host 
in List I. 

The incidence of helminths in birds dissected by us 
and by the late T.H. Johnston is noted and brief 
comments made on the birds and on the helminths, 
A complete reference list is also given, 








LIST 1. Birds in Taxonomic Order, and 

their Parasites . 223 

LIST 2. Helminths in Taxonomic Order, 

and their Hosts 284 

LIST 3. Index to Families of Birds . - 305 

LIST 4. Index to Species of Birds 306 

LIST 5. Index to Helminths 310 

REFERENCES. . . . 320 


No complete list of helminths from Australian 
birds— not even a complete list of any major group of 
helminths— has been published for over forty years. 
During this time a great deal of research has been done 
on the taxonomy of helminths from Australian birds, 
much of it by the present authors, and it is considered 
timely to draw all these records together. 
Comprehensive lists published earlier than 1923 were 
made by T.H. Johnston (1910, 1912b, 1918) and J.B. 
Cleland (1922). Partial lists dealing with the parasites 

in restricted geographical areas or with one group of 
parasites were published bv Bancroft (1889), Cleland 
and T.H. Johnston (1910, 1912), T.H Johnston (1916), 
Brcinl (1913), Nicoll (I^Ua.b), and T.H. Johnston and 
Deland (1929). In 1939 MR. Young published a list of 
helminth parasites of Australia. Since then, restricted 
lists have been issued by Mackcrras (1962) on filarial 
nematodes of Australian vertebrates and by Munday 
and Green (1972) on all helminth parasites of 
Tasmanian vertebrates. 

Without these earlier works the compilation of the 
present list would have been a formidable task, The 
present authors are indebted to them and to the host 
and parasite files started by T.H. Johnston and now 
continued as part of the Australian Helminthological 
Collection (at present housed in the South Australian 

Very little serious collecting Of helminths from birds 
has taken place in Australia. Most known helminths 
have been found incidental to bird collection. Museum 
ornithologists who collect birds for taxonomic 
purposes have usually no need to keep the carcases 
when the skins are removed, so that unless a 
helminthologist can arrange to obtain a carcase from 
a bird collector, much valuable helminthological 
material may be lost. We have been fortunate in having 
such liaison with successive ornithologists (H.T. 
Condon and S.A, Parker) at the South Australian 
Museum, and with officers of the (then) Animal 
Industry and Agriculture Branch of the Northern 
Territory Administration, who undertook extra work 
in labelling and transmitting to us the frozen, skinned 
torsos of birds which they had collected. 

This list of helminths is concerned only with those 
from native Australian birds; the parasites of 
introduced birds, caged or feral, have not been 
included. Nor have birds taken on Macquarie Island 
or in the Australian Antarctic Territory been included. 
We have found no case of interchange of helminth 
parasites between introduced and native birds dissected 
by ourselves except when the latter are in captivity. 
Syngamus trachea has been recorded by others from 
uncaged birds, as noted in the text. 


Information for this checklist was obtained from all 
major Museums and collections in Australia and from 
the British Museum (Natural History). We are grateful 
for all the trouble taken by the staff of these institutions 
in sending details of relevant material and, in some 
cases, specimens. 

Atu'usL 198& 


REC. S. AUST. MLIS. 19(15): 2f£323 

.4tfS*w/, /?S6 

We wish to acknowledge special help from Mr Shane 
Parker with the nomenclature of birds, Dr Gordon 
Gross for advice on the arrangement of the manuscript, 
and to many helminthologists in Australian universities 
for help in obtaining specimens and with references* 
and identifications. 

Our warm thanks go also to colleagues in the 
Zoology Department of the University of Adelaide and 
the South Australian Museum, and to friends in various 
places, who have responded with patience and tolerance 
to our frequent inquiries about Australian birds and 
their helminth inhabitants. 


This checklist has been compiled from ail published 
records up to the end of 1983, which are known to us, 
and from unpublished material, lists of which have 
been made available to us, held in Museums and other 
institutions. Published work which is merely a 
repetition of an earlier record is not included, Some 
of the material listed is not fully identified. This applies 
particularly to cestodes, as less work has been done in 
Australia on these parasites than on other helminths; 
and because none of the authors can claim expertise 
in this field. All specimens* whether identified or not, 
have been included, not only to show the range of 
parasitism, but also in the hope that interested 
taxonomists may wish to examine the specimens, We 
have not ourselves seen all the material listed, and have 
examined only that in the Australian Helminthologieal 
Collection, that described in our own papers, and some 
of that in the Veterinary School, University of 
Melbourne and in the Commonwealth Institute of 
Health, University of Sydney. 

In this work the information i.s arranged in the 
following sections: 

List I. Bird species arranged systematically and 
numbered consecutively, with the helminths 
from each listed in taxonomic order under it; 
the locality and authority are given for each 
record. Synonymy is indicated where necessary. 

List 2. Helminths arranged systematically, with bird 
hosts under each, and with numerical 
references to List I. 

List 3. Families of birds listed, in alphabetic order, with 
page numbers (in this paper). 

List 4. Bird species listed alphabetically, with numbers 
referring to List L 

List 5. Helminth species arranged alphabetically in 
each of the four helminth groups, with 
numerical references to List 1. Synonyms 
named in List I are included in alphabetic 


J.B. Cleland (1922) gave a list of parasites from 
Australian birds. He included, as well as a list of earlier 
records, a section on birds examined by himself and 
the parasites found in them. The helminths in this 
section were seldom identified further than to one of 
the four helminth groups. The present whereabouts of 
many of these specimens is not known, but some have 

been described since, and some are in the Australian 
Helminthologieal Collection and arc noted as such in 
List 1. 

Abbreviations used in the text; 
Authors whose names appear frequently arc referred 
to, where appropriate, by initials, as follows: 
LMA I.. Madeline Angel 
S.TE Stanley J. Edmonds 
SJJ Stephen J, Johnston 
THJ T. Harvey Johnston 
PMM Patricia M. Mawson 
The four major groups of helminths are referred to 
by their initials: 
A Acanthocephala 
C Cestoda 
N Nematoda 
T Trematoda 

Museums and other Institutions which are referred 
to as having bird helminths in their collections are 
abbreviated as follows: 

AHC Australian Helminthologieal Collection, at 
present housed in the South Australian 
Museum, Adelaide, South Australia 
AM Australian Museum, Sydney, New South 

BM(NH) British Museum {Natural History), 

London, England 

CIH Commonwealth Institute of 

Helminthology, St Albans, Herts, England 

CIHUS Commonwealth Institute of Health, 

University of Sydney, New South Wales 

DWRR Division of Wildlife & Rangelands 

Research (C.S.I. R.O.), Canberra, A.C.T, 
NMV (Formerly National) Museum of Victoria, 

Melbourne, Victoria 
QM Queensland Museum, Brisbane, 

SAM South Australian Museum, Adelaide, 

South Australia 
UMVS University of Melbourne, Victoria, School 

of Veterinary Science 
UQDP University of Queensland, Brisbane, 

Department of Parasitology 
WAM Western Australian Museum, Perth, 
Western Australia 

Localities from which birds and parasites are 
recorded are given as the Stale in which they lie. though 
more detail is usually available in the references cited. 
State names are abbreviated as follows; 

ACT Australian Capital Territory 

FA Eastern Australia 

NSW New South Wales, including Lord Howe Island 

NT Northern Territory 

Qld Queensland, including islands of the Great 
Barrier Reef 

SA South Australia, including Kangaroo 1. and 
Pearson 1. 

Tas Tasmania, including the Bass Strait Islands 
(King I. & Flinders 1.) 

Vic Victoria 

WA Western Australia 

If a host is known to have been captive (cage, aviary, 
etc.) the record is marked 'cage'. 




Bird nomenclature, both English and scientific, 
and the order in which the names arc listed, follow that 
of Schodde et ai (1978), except for certain amendments 
made since 1978. 

The birds in the host-parasite list have been 
numbered 1-443. These numbers do not refer to any 
previously published list of Australian birds, but are 
given to facilitate cross-reference from oilier lists in this 
work to The first, which contains greater reference 

In the first list, birds which migrate regularly to and 
from other countries are noted as •migratory*, the term 
here not including those which migrate within 
Australia, or ihose which occur outside Australia but 
arc not regular migrants. If a bird occurs only in 
Australia ir is noted as 'endemic*. We have not included 
helminths taken from migratory birds, or from those 
which occur naturally in othei countries, if they were 
collected outside their Australian range. However, we 
have included records from birds of distinctly 
Australian origin in captivity in other countries. Any 
uYord from a bird known to have been in captivity, 
in Australia or elsewhere, is marked '(cage)*. As 
mentioned earlier, helminths from birds introduced into 
Australia since the advent of the white man have not 
been included. It is hoped that this omission may be 
rectified later. 

The Cattle Egrcl, Anieola ibi$\ was not actually 
introduced into Australia by man, It is thought to have 
followed the introduced Water Buffalo, after this had 
become established in northern Australia. The Cattle 
Egret was first noted by ornithologists to be present 
in the late 1340s, and has now extended its range to 
southern Australia. 

The legends (numbers and letters) which appear 
directly under the name of some birds in List I refer 
to autopsies undertaken by ourselves or by Professor 
T,H, Johnston Careful records of these dissections 
were kept in the Zoology Department, University of 
Adelaide, by Johnston from 1922 to his death in 1951, 
and since then by Angel and Mawson. In these legends, 
the figures indicate the number of birds 
examined /number found with helminths, and the 
letters refer to the group of helminth, the number after 
each tetter being the number of birds infested with thai 

Seven hundred and six species of native Australian 
birds, representing 82 bird families, are listed by 
Schodde et al. 1 1978) Helminths are known to occur 
in 384 (54%) of these birds; at least 55 (S'Vo) other 
species have been examined but have not yielded 
helminths. This means that about 40% of the 
Australian birds arc unexplored territory for internal 
parasites. Bird families represented in Australia from 
which no helminths have been recorded arc (with the 
number of Australian species in brackets) Pandionidae 
(!); Rostratulidae (1); Phalaropidae (3); Stereorarudae 
(5); Oposirtidae (I); Nectarinidae (I); Dicaeidae (1); 
AhrK-homitnidac (2); and Sturnidae (1). In List 3 the 
number of Australian bird species ^n eaeh family is 
noted after the family name, together with the number 
from which helminths have been recorded. 

Before the studies of Rowley (1967, 1970) the 
distribution and taxonomy of Australian corvids were 
poorly known or understood, Indeed it was not umil 
these studies were made that Corvus metlori (Little 
Raven) and C. tasmamensis (Forest Raven) were 
recognised as species distinct from C corortoides 
(Australian Raven), For a substantial number of early 
records, the identity of the corvid hosts can be 
established by recourse to the original (museum) skim 
or reasonably assumed from the locality involved. For 
othei old records, however, where host specimens were 
not retained and where the provenance falls within a 
zone of overlap between (he now-reeognised species, 
the identity of the corvid host will probably always 
remain uncertain. Records from such uncertain origins 
are marked ■ ? host sp;\ 


[n general, the helminths from Australian birds 
resemble those from other parts of the world. There 
are veiy few genera which are (united to Australian 
birds, although many species and some sub-species arc 
so restricted. Some helminth species arc found in 
related birds in many pans of the world, such overlap 
may become more evident with greater knowledge of 
the \arious genera. 

Helminths of the gut are rare in free-ranging 
psittacifoTm birds in Australia. Of about 160 such birds 
dissected by us, cestodes were present in 5, nematodes 
(minute larvae) in 2, and trematodes and 
aeanthocephalans in none (PMM, 1985, p, 190) 
However, the trcmatode Platynasomum fjroxillicens 
and the nematodes Ascandia cohttnbae and A. 
nlafyceri have been taken from native birds which have 
been held in captivity in Australia or in other countries. 
Two possible reasons for this may be that Ihe caged 
bird is more vulnerable to infection (and many of the 
infestations were very heavy and were said to have 
caused the death of the host), or that it is exposed to 
the eggs or larvae of parasites through proximity to 
other (? introduced) caged birds or to domestic .stock 
such as pigeons, Ascarldia pfaiyceri has recently been 
found in a feral colony of Agapornis roseicollis near 
Cowell, SA (Coll.S, Paiker, idem. PMM, in AHO. 

Birds dissected by ourselves had often been frozen 
before we received them, a treatment particularly 
damaging to Hat worms. However, rather than 
disregard poor specimens we have listed them to record 
their presence, and given what classification is possible- 

tn this work we have not referred to sub-species, 
either of helminths or hosts. 


The classification and nomenclature followed hctc 
arc for the most part those of Yamaguti (1971). The 
main exception is in the Strigcidae, in which we folio* 
Dubois (1938 et seq.). Entries in the checklist arc placed 
according to the order in which families are placed by 
Yamaguti (1971,p. 475) Families of Digenca from fish 
also follow the order used by Yamaguti (1971* p. 19) 
(Cysts of these trematodes are often recorded from the 
aJunemary canals of birds). 


RK . 5, AUST. MUS. 19(15); 219-3Z5 

.4w^/y>7. /0*6 

Since the number and arrangement of collar spines 
is important in the identification of echinostomes, they 
have been recorded fOT incompletely indentified 
specimens of this group in one of the following ways, 
as appropriate: 
Unidentified echinostome; [spines (35(5))] indicates 

that the total number of spines is 35, including 5 

corner spines on each side. 
Hehinostomatidae: [spines missingj. 
lichmostomatmac: [spines? (35(5))]: total number ?35, 

including 5 corner spines on each side, 
in Echinochasmus sp.: [spines 20]; total number 20, 

no corner spines. 
In Patagifer sp.: [spines 2 x (30(4))]* total number 60. 

30 on each side including 4 corner spines. 
In Patagifer sp.: [spines 2 x ? (29(4))l: total number 

?5S, including 4 corner spines on each side. 


For the most part the names used for cestodes follow 
the classification adopted by Yamaguti (1959). Other 
more recent works consulted in special cases were Baer 
and Bona (1960) and Bona (1975) for cestodes of 
Ciconiformes, Palmer (1981) for cestodes of the Black 
Swan, Cygnus atraius. and Schmidt (1972) for some 
cestodes from honeyeaters (Meliphagidae) and parrots. 
None of the present authors is well versed in cestode 
taxonomy; cestode lists here are the responsibility of 

It may be noted here that RA. Maplcstone deposited 
four lots of cestodes in the Australian Institute of 
Tropical Medicine (now the Commonwealth Institute 
of Health* University of Sydney) under names which 
apparently have never been published. All are labelled 
as 4h new species, Maplcstone" The names are "Shipleya 
lobixanellus'- and *%fonopyiidium fobivanelti" both 
from "Yuneilus miles') f Hymenulepis variabilis" from 
Cygnus atrari4s; and "H. eurysiomi" from Eurystomus 
orientalis. The explanation winch occurs to us is that 
Maplcstone examined this material while he was in 
Queensland, decided that it belonged to these four new 
species, and deposited paratypes in what is now the 
CIHUS Collection before returning to England. After 
his return, Maplestone published a number of papers, 
some with Southwell, on Australian cestodes, from the 
same hosts and localities but with different scientific 
names; one such species is Monopylidium 
macracanthum Maplestone and Southwell, 1923, which 
may be the unpublished "M, lobnanelli^ In one of the 
papers he identified Hymenolepis lanceolata (Bloch, 
17821 from Cygnus atratus; the CIHUS material 
labelled *'H. variabilis Maplestone' may belong to this 

In the Australian Helminthological Collection there 
are slides of the first two of these unpublished species, 
presumably given to T.H Johnston by Maplestone when 
they were both in Queensland. The AHC material 
labelled "Shipleya lobivanellus Maplestone' has been 
examined by Schmidt (1972) who identified it as hifula 
burhini Burt, 1939, 


Nomenclature in this section follows on the whole 
that adopted by Anderson. Chabaud and Wiltmotl, 
1974-1982, (OH keys to the nematode parasites of 
vertebrates) as far as these keys are completed. Some 
points to be noted are; 

Anisakis diometieae (Linstow, 1888) is recorded from 
a number of procellariform birds. The validity of this 
species has recently been upheld (Mawson 1983, p. 
247). It is the only species of the genus Anisakis known 
from birds, but is commonly found in albatrosses and 
petrels along the Australian coast. Linstow-' described 
it, as Ascaris diomedeae % from material collected by 
the Challenger Expedition in the northern Pacific 
Ocean, from Diotnedea brachyura, 

Cartfracvecum spicuhgerum (Rudolph*. 1809), 
commonly recorded from warerhirds was considered 
by Hartwich (1964) to be a junior synonym ol C 
microcephalum (Rudolphi, 1809), and Hanwich 
suggested that the specimens hitherto allotted to 
C spiculigerum belong cither to C mierocephalum or 
to C rudolphi Hartwich. 1964. However, in this 
checklist the name C spiculi^rum is retained, pending 
Turther review of the Australian species of 
Contracaecum, Hartwich also suggested that C 
nwgnipapitlatum Johnston and Mawson, 1941, is a 
junior synonym of C variegawm (Rudolphi, 1809) but 
the two species appear to us to be quite distinct. 

Three different species of Schistogendria have been 
examined from various birds, but have not been 
described. They are referred to by numbers; voucher 
specimens are in the AHC. 

Cyrnea spiralis Mawson, 1968d t is now considered 
likely to belong to, or near, Microhadjelia Jogis, 1968, 
and is referred to in this work as ? Microhadjelia 


The system of classification of acanthocephalans 
used in this paper is that of Yamaguti (1963). However, 
most of the changes in nomenclature made since 1963, 
especially those of Schmidt (1983) have been adopted. 
No new genus of Acanthocephala from Australian 
birds ruts so far been recorded. 

The embryonated eggs of an acathocephalan are 
shed with the faeces of the host. The first intermediate 
host in the life cycle of the parasite is thought to be 
always an arthropod, which is infected by eating the 
eggs. Sometimes a second intermediate host (often 
called a paratenic host or "l*hote d'attente") is 
necessary. A final or definitive host is infected when 
it ingests an infected intermediate host. Consequently 
adult acanthocephalans in birds occut \u those which 
are partly or wholly carnivorous, This possibly explains 
why no acanthocephalans have so far been collected 
from the Emu, Dmntaius rtovaehoitandiue, or from 
psinaciform birds, which are principally seed eaters 

The life cycles of Australian avian acanthocephalans 
are poorly known and none have been determined 
experimentally. What information about life cycles is 
available has been obtained by a comparison of the size 
and armature of the introverts of larval and adult 



forms. Using such a method TH. Johnston and 
Edmonds (1949) considered that one of (he larva! stages 
of Porrorchis (= Gordiorhynchus) hylae (Johnston, 
1912), the adult stages of which occur in the birds 
Podargus strigoides and Centropus phasianinus, is 
found as a cyst in the mesenteries and muscles of a 
number of frogs. The frog is infected, it is assumed, 
by eating an infected crustacean or insect. Some light 
has been thrown on the life cycles of two other avian 
Acanthocephala. The adult form of Polymorphic 
biziuraeT.H. Johnston and Edmonds (1948) occurs in 
the small intestine of the Musk Duck, Biziura lobata. 
T.H. Johnston and Edmonds (1948) considered that the 
freshwater yabbie, Cherax destructor, is an intermediate 
host of the parasite. One of the commonest 
acanthocephalans known to dissectors of birds in 
Australia is a small cyst, 2-3 mm long and shaped like 
a "grain of rice", which occurs in the subcutaneous 
tissues of the neck and thorax of a large number of 
passerine birds. It was called Echinorhynchus 
pomatosiomi by I.H.Johnston and Cleland (1912). 
Schmidt (1983) has found that the introvert of E. 
pomatosiomi matches that of an acanthocephalan 
found in the intestine of dingos and feral cats in 
Australia. From this information he has concluded that 
the cyst known as E . pomatostomi is an early stage of 
the acanthocephalan found in dingos and feral cats. 
Since I he adult form is an Oncicola and not an 
Echinorhynchus, the parasite (including the cyst) is now 
Onacola pomatostomi (T.H. Johnston and Cleland, 
1912). The carnivores are infected by ingesting the cysts 
in their bird prey. The first host is probably a 
scavenging insect, though at present there is no evidence 
for this. 


Explanation of the symbols used in this section are 
given in "Comments on bird hosts" on p. 221. 

1. Dromaius novachallMdiae (Latham) 

Emu (endemic) 
l/l : C 

T. Philophlha/mus sp t 

C. Cotugnia collini Fuhrmann, 1909 
EA: Fuhrmann, 1909, p. II G 
Cotugnia sp. 

Raitiietina uustralis (Krabbe, 1869) 
syn. Davainea australis (Krabbe, 1869) 
" NSW: THJ, 1909c, p.xxix 
NSW, WA: THJ. 1910, p,86 
Qld: THJ, 1916, p.45 
syn. Taenia australis Krabbe, 1869 
Copenhagen (cage): Krabbe, 1869, p.343 
Railitettna sp. 



Qld, NSW, Vic, SA, WA: AHC 
N. Dromaeostrongylus bicuspis Lubimov, 1933 
Moscow (cage): Lubimov, 1933, p.173 
Vic: Durette-Desset, 1979, p.1015 
NSW, Vic: AHC 
Trichostrongvlus tenuis (Mehlis, 1846, in Creplin, 
Aust: Arundel, 1982, pers. comm. 


2. Casuarius casuarius (Linne) 
Southern Cassowary 

C, Unidentified 

Old: Macgilltvray, 1917, p.80 


3. Podiceps cristatus (Linne) 

Great Crested Grebe 

5/5 : Tl C4 N4 

T. Schistosomal idae, unidentified 
Pe/asiger australis TH. Johnston & Angel, 1941b 

Diptostomum podicipinum Kozieka & 
Niewiadomska, 1960 
SA: Dubois & LMA, 1972, p.208 
Schwartzitrema pandubi (Pande, 1939) 

SA: Dubois & LMA, 1972, p.205 (cysts) 
C. Unidentified 
Krcfft (1873, pp.216,217) recorded Taenia paradoxa and 
T novaehollandiae from the "Little Grebe (Podiceps 
australis)". P. australis is now placed as a synonym of 
the Crested Grebe, (R cristatus), and it is considered 
that Krefft's records refer to Tachybaptus 
novaehollandiae, which was then called the Little Grebe 
N. Capillaria sp. 

SA: THJ & PMM, 1949, p.64 
Contracaecum podicipitis T,H. Johnston & 
Mawson, 1949 
SA: THJ & PMM, 1949, p.67 
NSW, Vic: AHC 
Tbtrameres gubanovi Shigin, 1957 

SA: PMM, 1979, p.180 
Svncuaria sp. 

ACT. PMM, 1982, p.2I 

4. Polioeephalus poliocephalus 

(Jardine & Selby) 

Hoary-headed Grebe (endemic) 

5/4 : T3 C3 N3 

T. Paramonostomurn caeci Smith & Hickman, 1983b 
Tas: Smith & Hickman, 1983b, p.86 
Cvclocoelum jaenschi TH. Johnston & Simpson, 


RFC. S. AUST. MUS. W5): 21W25 

4ugiU£ /JWto 

SA: THJ & Simpson, 1940b, p.273 
Echinostoma sp. [spines (37(4))] 

Patagifer sp. [spines 2x (26-27(3-4))] 

Qld: AHC 
Petasiger australis T.H. Johnston & Angel, 1941b 

SA: THJ & LMA, 1941b, p.285 
Echinostomatidae, unidentified 

NSW: Bradley, 1926, p.575; 1927, p.675 
Psilochasmus oxyurus (Creplin, 1825) 

Tas: S.J. Smith, 1981, p.181 
Psilostomum sp. A, S.J. Smith, 1981 

Tas: S.J. Smith, 1981, p.201 
Psilostomum sp. B, S.J. Smith, 1981 

Tas: SJ. Smith, 1981, p.214 
Atriophallophorus coxiellae S.J. Smith, 1974 

Tas: S.J. Smith, 1981, p.151 
Levinseniella tusmaniue (S.J. Smith, 1974) 

Tas: S.J. Smith, 1981, p. 107 
Mahtrema calvertense S.J. Smith, 1974 

Tas: S.J. Smith, 1981, p.64 
Pachytrema sp. 

Tas: S.J. Smith, pers. comm., 1979 
Sehwartzitrema pandubi (Pande, 1939) 

SA: Dubois & LMA, 1972, p.205 (cysts) 
? Cephalogonimus sp. 

C. Dioecocestus sp. 

Qld: BM(NH) 
Gyrocoelia sp. 

Aust: BM(NH) 

NSW: Bradley, 1926, p.572, 1927, p.675 

NSW: Cleland, 1922, p.105 

Qld, NSW, SA: AHC 
N. Capiliaha sp. 

Contracaecum praesiriatum Monnig, 1923 

Qld: C1HUS, BM(NH) 
Streptocara recta (Linstow, 1879) 

SA: THJ & PMM, 1941e, p.16 

? !oc: Bradley, 1927, p.675 

5. Tachybaptus navaehoMandiae 


Australian Grebe 

13/10 : T8 CIO N8 

T. Cyclocoelum jaenschi T.H. Johnston & Simpson, 

SA: THJ & Simpson, 1940b, p.273 
Petasiger australis T.H. Johnston & Angel, 1941b 

SA: THJ & LMA, 1941b, p.285 

Echinostomatidae, very immature, [spines ? (37)] 

SA: discarded 
Mahtrema oocystum (labour, 1907) 

Qld: Deblock & Pearson, 1968b, p.459 
Microphatlidae, unidentified 

SA: (believed lost) 
Schwartzitrema pandubi (Pande, 1939) 

SA: Dubois & LMA, 1972, p.205 (cysts) 

C. Dilepididae, unideniified 
Dioecocestus novaehollandiae (Krefft, 1873) 
syn. Taenia novaehollandiae Krefft, 1873 

NSW: Krefft, 1873, p.216 
syn. Taenia paradoxa Krefft, 1873 
NSW: Krefft, 1873, p.217 

Qld, NSW, SA: AHC 
N. Contracaecum podicipitis T.H. Johnston & 
Mawson, 1949 
SA: THJ & PMM, 1949, p.67 
Tetrameres gubanovi Shigin, 1957 

NSW: PMM, 1979, p.180 
Skrjabinoclava horrida (Rudolphi, 1809) 

Qld: AHC 
Streptocara recta (Linstow, 1879) 

SA: THJ & PMM, 1941e, p.260; PMM, 1955, 

Streptocara sp. 

Svncuaria sp. 


6. Podicipedidae 

Unidentified grebe 

C. Unidentified 

Qld: AHC 
N. Capilluria sp. 

Qld: AHC 


1. Aptenodytes patagonicus Miller 
King Penguin 

C. Tetrabothriidae, unidentified 
Tas: AHC 

8. Eudyptes chrysocome (Forster) 

Rockhopper Penguin 

N. Anisakidae 

Tas: AHC (larva) 

9. Eudyples pachyrhynchus Gray 

Fiordland Penguin 

N. Contracaecum sp. 
Tas: AHC 

10. Eudyptes schlegeH Finsch 

Royal Penguin 

C. Unidentified 
Tas: AHC 

11. Eudyptula minor (Forster) 

Little Penguin 
4/4 : T2 C4 N4 A2 


T. Mawsonotrema eudyptulae Angel, 1973 

SA: LMA, 1973, p.857 

Vic: UMVS 

Tas: AHC 
Renicola sp. 

Tas: AHC 
Galuctosomum angelae Pearson, 1973 

SA: Pearson, 1973, p.361 
C. Tetrahothrius lutzi Parona, 1901 

Tas: Prudhoc, 1969, p.185 
Tetrahothrius sp. 

NSW, Vic, SA: AHC 
Tetrabothriidac, unidentified 

Vic: AHC 

SA: Cleland, 1922, p.105 

NSW, Vic, Tas, SA: AHC 

N. Anisakis sp. 

NSW, SA: THJ & PMM, 1942a, p.94 
Contracaecum spiculigerum (Rudolphi, 1809) 

Vic, Tas, SA: AHC 
Contracaecum eudyptulae T.H. Johnston & 
Mawson, 1942a 
NSW, Vic, Tas, SA, WA: THJ & PMM, 1942a, 
syn. unidentified nematode 
SA: Cleland, 1922, p.107 
Fiiarioidea, unidentified 

SA: AHC (from heart) 
Vic: UMVS 
A. Corynosoma sp. 


12, Diomedea exuians Linne 

Wandering Albatross (migratory) 
1/1 : C N 

Stegophorus sp. 

13. Diomedea melanophrys Temminek 

Black-browed Albatross 

1/1 : C N 

C. Tetrahothrius sp. 

NSW: THJ, 1912b, p.106 
Tas: AHC 
Cyclophyllidea, unidentified 

Tas: Munday & Green, 1972, p.4 

N. Capillaria convolutor Fourment, 1885 
SA: THJ & PMM, 1945a, p.151 
Anisakis diomedeae (Linstow, 1888) 

NSW, SA: THJ & PMM, 1942d, p.67 
syn. Stomachus sp. 

Tas: Munday & Green, 1972, p.9 
Contracaecum pelagicum T.H. Johnston & 
Mawson, 1942d 
NSW, SA: THJ & PMM, 1942d, p.67 
Anisakidae, unidentified 

Tas: Munday & Green, 1972, p.9 
Seuratia shipleyi (Stossich, 1900) 

SA: THJ & PMM, 1942d, p.69 
Stegophorus diomedeae (T.H. Johnston & Mawson, 
syn. Paryseria diomedeae T.H. Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942d f p.67 

14. Diomedea butteri Rothschild 
Buller's Albatross 

N. Seuratia shipleyi (Stossich, 1900) 

C, Tetrahothrius diomedeae (Fuhrmann in Shipley, 
syn. Prosthecocotvle diomedeae Fuhrmann in 
Shipley, 1900 
'Pacific Ocean, NE of Austr. 7 : 
Fuhrmann, in Shipley, 1900, p.557 
Tetrahothrius sp. 

NSW: THJ, 1912b, p.106 

N. Anisakis diomedeae (Linstow, 1888) 

NSW, Tas: THJ & PMM, 1942d } p.6 
syn. Contracaecum diomedeae (Linstow, 1888) 
Tas: THJ, 1938, p.15 
Contracaecum sp. 

SA: AHC (immature) 
Seuratia shipleyi (Stossich. 1900) 

Stegophorus diomedeae (T.H. Johnston & Mawson, 
syn. Paryseria diomedeae T.H. Johnston & 
Mawson, 1942d 
NSW: THJ & PMM, 1942d, p.69 

15. Diomedea chrysostoma Forster 

Grev-headed Albatross 
4/4 : CI N4 

C. Unidentified 

N. Anisakis diomedeae (Linstow, 1888) 
SA: THJ & PMM, 1942d, p.67 
lis: AHC 
syn, Stomachus sp. 
Tas: Munday & Green, 1972, p.9 
Seuratia shipleyi (Stossich, 1900) 

SA: THJ & PMM, 1952, p.33 
Stegophorus diomedeae (T.H. Johnston & Mawson, 
syn. Paryseria diomedeae T.H. Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942d, p.69; 1952, p.33 
Stegophorus macronectes (T.H. Johnston & 
Mawson, 1942d) 
syn. Paryseria macronectes T.H. Johnston &. 
Mawson, 1942d 
SA: THJ & PMM, 1942d, p.70 

REC. S. AUST. MUS. 19(15): 219-325 

August, 1986 

Dlomedenema diomedeae T.H. Johnston & 
Mawson, 1952 
SA: THJ & PMM, 1952, p.32 

16. Diomedea chlororhynchos Gmelin 

Yellow-nosed Albatross 
3/3 : C3 N3 

C. Tetrabothrius sp. 
N. Anisakis diomedeae (Linstow, 1888) 
SA: THJ & PMM, 1942d, p.67 
Contracaecum pelagicum T.H. Johnston & 
Mawson, 1942d 
NSW: THJ & PMM, 1942d, p.67 
Tetrameres certa (Leidy 1886) 
syn. Tetrameres diomedeae T.H. Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942d, p.67 
Seuratia shipleyi (Stossich, 1900) 
SA: THJ & PMM, 1942d, p.67 

17. Diomedea cauta Gould 

Shy Albatross 

1/1 : C N 

C. Unidentified 

SA: not kept 
Tas: AHC 
N. Anisakis diomedeae (Linstow, 1888) 
SA: THJ & PMM, 1942d, p.67 
Vic: UMVS 
Tas: AHC 
Contracaecum magnico/lare T. H . Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942d, p.67 
Tetrameres certa (Leidy, 1886) 
syn. Tetrameres diomedeae T.H. Johnston & 
Mawson, 1942d 
Tas: AHC 
Stegophorus diomedeae T.H. Johnston & Mawson, 

Tas: AHC 

Tas: AHC 


T.H. Johnson & 

C. Tetrabothrius sp. 


Qld, SA: AHC 
N. Capillaria convolutor (Fourment, 1885) 
Anisakis diomedeae (Linstow, 1888) 
SA: THJ & PMM, 1942d, p.67 
Phocascaris sp. 

SA: THJ & PMM, 1942d, p.68 
Seuratia shipleyi (Stossich, 1900) 
SA: THJ & PMM, 1942d, p.69 
Stegophorus macronectes (T.H. Johnston & 
Mawson, 1942d) 
syn. Paryseria macronectes T.H. Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942d, p.70 
A. Unidentified 

SA: not kept 

20. Fulmarus glacialoides (Smith) 
Southern Fulmar 

C. Unidentified 

N. Seuratia shipleyi (Stossich, 1900) 


21. Daption capense (Linne) 
Cape Petrel 
2/2 : CI N2 

C Unidentified 

N. Anisakis diomedeae (Linstow, 1888) 

SA: THJ & PMM, 1942d, p.67 
Seuratia shipleyi (Stossich, 1900) 

SA: THJ & PMM, 1942d, p.69 
Stegophorus pachyptilae (T.H. Johnston & 

Mawson, 1942d) 

Spirurida, unidentified 

Tas: Munday & Green, 1972, p.10 

22. Pterodroma macroptera (Smith) 
Great-winged Petrel 

18. Phoebetha palpebrata (Forster) 
Light-mantled Sooty Albatross 

C. Tetrabothrius sp. 

N. Contracaecum sp. 

Vic: AHC 
Seuratia shipleyi (Stossich, 1900) 


N. Anisakis sp. 

Tas: AHC 
syn. Stomachus sp. 
Tas: Munday & Green, 1972, p.9 
Seuratia shipleyi (Stossich, 1900) 

Tas: Munday & Green, 1972, p.ll 
Tas: AHC 
Spirurida, unidentified 

Tas: Munday & Green, 1972, p.9 

19. Macronectes giganteus (Gmelin) 

Southern Giant Petrel 

10/9 : C2 N9 Al 

23. Pterodroma iessonii (Garnot) 

White-headed Petrel 

3/3 : CI N3 


C. Unidentified 

N. Anisakis sp. 

SA: THJ & PMM, 1942d, p.67 
Seuratia shipleyi (Stossich, 1900) 

Stegophorus macronectes (T.H. Johnston & 
Mawson, 1942d) 

24. Pterodroma brevirostris (Lesson) 

Kerguelen Petrel 
1/1 : N 

C. Tetrabothrius sp. 

N. Seuratia shipleyi (Stossich, 1900) 
Stegophorus macronectes (T.H. 
Mawson, 1942d) 

Johnston & 

25. Pachyptila vittata (Forster) 

Broad-billed Prion 
6/5 : C2 N5 

C. Tetrabothrius sp. 
N. Skrjabinociava sp. 
Tas: AHC 
Seuratia shipleyi (Stossich, 1900) 

SA: THJ Sl PMM, 1942d, p.69 
Stegophorus pachyptilae (T.H. Johnston & 
Mawson, 1942d) 
syn. Paryseria pachyptilae T.H. Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942d, p.70 

26. Pachyptila salvini (Mathews) 
Lesser Broad-billed Prion 

5/1 : N 

N. Seuratia shipleyi (Stossich, 1900) 
Stegophorus pachyptilae (T.H. Johnston & 
Mawson, 1942d) 

27. Pachyptila desolata (Gmelin) 

Antarctic Prion 
11/4 : N 

N. Anisakis sp., ? diomedeae (Linstow, 1888) 
SA: THJ & PMM, 1942c, p.184 
Stegophorus pachyptilae (T.H. Johnston & 
Mawson, I942d) 
syn. Paryseria pachyptilae T.H. Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942c, p.184 

28. Pachyptila belcheri (Mathews) 

Slender-billed Prion 

2/2 : N 

N. Anisakis sp. 

SA: AHC (immature) 
Seuratia shipleyi (Stossich, 1900) 

29. Pachyptila turtur (Kuhl) 
Fairy Prion 
4/4 : C3 N4 

C. Unidentified 

N. Skrjabinociava sp. 
Tas: AHC 
Seuratia shipleyi (Stossich, 1900) 

Stegophorus pachyptilae (T.H. Johnston & 
Mawson, 1942d) 
A. Plagiorhynchus sp. 
Tas: AHC 

30. Procellaria cincrea Gmelin 

Grey Petrel 

C. Unidentified 


31. Puffinus pacificus (Gmelin) 
Wedge-tailed Shearwater 

2/1 : N 

T. Galactosomum renincolum Pearson, 1973 
Qld: Pearson, 1973, p.409 

C. Unidentified 

NSW: Cleland, 1922, p.105 
N. Seuratia shipleyi (Stossich, 1900) 

Qld: AHC 

32. Puffinus griseus (Gmelin) 
Sooty Shearwater 

C. Tetrabothrius sp. 

NSW, Vic: AHC 

N. Contracaecum magnicollare T.H. Johnston & 
Mawson, 1941c 
Vic: AHC 
Stegophorus pachyptilae (T.H. Johnston & 
Mawson, 1942d) 
syn. Paryseria pachyptilae T.H. Johnston & 
Mawson, 1942d 
SA: THJ & PMM, 1942d, p.70 

33. Puffinus tenuirostris (Ternminck) 

Short-tailed Shearwater (migratory) 

12/8 : C3 N7 


KEC. S. AUST. MUS. 19(15): 219-325 

August, 1986 

T. Renicola sp. 

Cryptocotvle sp. 
Tas: AHC 

C. Tetrabothriidae, unidentified 
Vic: AHC 

Tas: Cleland, 1922, p.105 (AHC) 
Tas: Munday & Green, 1972, p.4 
N. Contracaecum magnicollare T.H. Johnston & 
Mawson, 1941c 
Vic: AHC 
Seuraiia shiplevi (Stossich, 1900) 
Vic, Tas, SA: AHC 
Bass Str: BM(NH) 
Stegophorus pachyptifae (T.H. Johnston & 
Mawson, 1942d) 
Tas, SA: AHC 
Stegophorus stellaepolaris (Parona, 1901) 
NSW, Bass Str: BM(NH) 
Tas: AHC 

34. Putfinus gavia (Forster) 

Fluttering Shearwater 

1/1 : C N 

T. Renicola sp. 

Tas: AHC 
C. Unidentified 

N. Unidentified 

SA: not kept (pieces only) 


35. Pclagodroma marina (Latham) 

White-faced Storm Petrel 

6/3 : N 

N. Seuraiia shiplevi (Stossich, 1900) 
Tas: AHC 
syn. Seuraiia marina T.H. Johnston & Mawson, 
Tas: THJ & PMM, 1941e, p.259; TH.I & PMM, 

1944, p.62 
SA: THJ & PMM, 1942d, p.69 
syn. "nematode" 

SA: Cleland, 1922, p.107 (AHC) 


36. Peleeanoides urinutrix (Gmelin) 

Common Diving Petrel 

2/2 : N 

N. Spirurida, unidentified 
Tas: AHC (larva) 


37. Pelevanus conspitillatus Temminck 

Australian Pelican 

12/12 : T12 C12 N12 Al 

T. Ornithobilharzia sp. 

Dendritobilharzia sp. 

Schistosomatinae, unidentified 

Old, SA: AHC 
Echinoehasmus pelecani T.H. Johnston & Simpson, 

SA: THJ & Simpson, 1944, p.113 

Qld: AHC 
Echinostomatidae [spines missing] 

Renicola sp. 

Qld: Pearson, 1979 (? 2 spp.) 

Qld: AHC 
Ascocoivie sp. 

Qld: AHC 
Haplorchis sprenti Pearson, 1964 

Qld: Pearson, 1964, p. 639 
syn. Haplorchis sp. 

Qld: Pearson, 1960, p.93 
Haplorchis paravanissimus Pearson & Ow-Yang, 

SA: Pearson & Ow-Yang, 1982, p.50 
Haplorchis vanissimus Africa, 1938 

SA: Pearson & Ow-Yang, 1982, p.50 
Procerovum varium Onji & Nishio, 1916 

Qld: Pearson, 1964, p.653 
syn. Procerovum sp. 

Qld: Pearson, 1960, p.93 
Stictodora caballeroi Martin, 1955 

Qld: Pearson, 1960, p.93 
Clinostomum sp. 

Qld: AHC 
Mesostephanus haliasturis Tubangui & Masilungan, 

SA: Dubois & LMA, 1972, p.214 
syn. M. minor Dubois & Pearson, 1965 

Qld: Dubois & Pearson, 1967, p.202 
Bolbophorus confusus (Krausc, 1914) 

Qld: Dubois & Pearson, 1965, p.95 

SA: Dubois & LMA, 1972, p.205 
Bolbophorus sp. 

SA: THJ & LMA, 1942a, p.59 
Diplostomum parvulum Dubois & Angel, 1972 

SA: Dubois & LMA, 1972, p.206 
Diploslomum sp. 

Qld: AHC 
Post hodiplostomum australe Dubois, 1937a 

SA: Dubois & LMA, 1972, p.212 
Schwartzitrema pandubi (Pande, 1939) 

SA: Dubois & LMA, 1972, p.205 (cysts) 
C. Hymenolepis murravensis T.H. Johnston & Clark, 

SA: THJ & Clark, 1948a, p.77 

*omitted by Yamaguti, 1959 


22 l J 

Hvmenolepis jaenschi T.H. Johnston & Clark, T. 
1948a* ' 

SA: THJ & Clark, 1948a, p.79 C. 

Qld: AHC 
Hvmenolepis ellisi T.H. Johnston & Clark, 1948af N 

SA: THJ & Clark, 1948a, p.8t 
Hvmenolepis sp. 


Qld, NSW, Tas, SA, WA: AHC 

N. Eusirongylides sp. 

Qld: CIHUS (posterior end o*, in gular pouch) 
Capillaria jaenschi T.H. Johnston & Mawson, 

1945b N 

SA: THJ & PMM, 1945b, p. 245 
Contracaecum clelandt'J.H. Johnston & Mawson, 
WA: THJ & PMM, 1941c, p.113 
Tas: AHC 
Contracaecum tnicropapillatum (Stossich, 1890) 
syn. Contracaecum hancrofti T.H. Johnston & 
Mawson, 1941c 
Qld, NSW, Vic, SA: THJ & PMM, 1941c, p.U3 
Contracaecum spiculigerum (Rudolphi, 1809) 
Vic: AHC 
syn. Ascaris spiculigerum Rudolphi, 1809 

NSW: THJ, 1912a, p.74 N. 

Tetrameres greeni Mawson, 1979 
Qld: PMM, 1979, p.180 
syn. Tetrameres pelecani T.H. Johnston & 
Mawson, I942e, sensu T.H. Johnston & 
Mawson, 1942c 
SA: THJ & PMM, 1942c, p.185 
Microietrameres pelecani ( T. H . Jo h nston & 
Mawson, 1942e) 
Qld: PMM, 1979, p.182 
syn. Teirameres pelecani T.H. Johnston & 
Mawson , 1942c nee 1942c 
SA: THJ & PMM, 1942e, pJ2 
Cosmocephalus jaenschi T.H. Johnston & Mawson, 
SA: THJ & PMM, 1942e, p.185 
Svnhimantus sirrv Khalil, 1931 
Qld, Vic: PMM, 1982, p.23 
syn. Dispharynx pelecani T.H. Johnston & 
Mawson, 1942c 
SA: THJ & PMM, 1942c, p.185 
Filarioidea, unidentified 
Qld: AHC (from eye) 
A. Potvmorphus hiziurae T.H. Johnston & Edmonds, 

Family StjUOAE 

38. Sula serrator (Gray) 

Australasian Gannet 
6/4 : Tl CI N4 

Galactosomum angelae Pearson, 1973 

SA: Pearson, 1973, p.361 
Tetrabothriidae, unidentified 

Contracaecum magnicollare T.H. Johnston 
Mawson, 1941c 

Tas: AHC 
Contracaecum sp. 


39. Sula daclylalra Lesson 
Masked Booby 


40. Sula sula (Linne) 

Red-footed Booby 

I/l : N 


N. Unidentified 

Qld: AHC 

41. Sula leucogaster (Boddaert) 
Brown Booby 

2/1 : N 

Seuratia shiplevi (Stossich, 1900) 
Qld, SA: AHC 


42. Anhinga melanogaster Pennant 


listed by Yamaguti, 

1959 in appendix to 

Mahtrema oocyst urn (Lebour, 1907) 

Qld: Deblock & Pearson, 1968b, p.459 
Clinostomum australiense S.J. Johnston, 1917 
Qld: SJJ, 1917, p.230 
Qld: THJ, 1942b, pl87 
Mesostephanus haliasturis Tubangui & Masilungan, 
syit A/, minor Dubois & Pearson, 1965 
Qld: Dubois & Pearson, 1967, p.202 
Diplosiomum auriculosum Dubois & Pearson, 1967 

Qld: Dubois & Pearson, 1967, p.193 
Schwarrzitrema novaehollandiae Dubois & 
Pearson, 1967 
Qld: Dubois & Pearson, 1967, p.187 
N. Eustrongvlides p/orinus T.H. Johnston & Mawson, 
Qld: THJ & PMM, 194Ie, p.256 
Contracaecum rodhaini (Gedoelst, 1916) 

Contracaecum sinulabiatum T.H. Johnston & 
Mawson, 1941c 
Qld: THJ & PMM, 1941c, p.U4 
syn. Ascaris spiculigerum Rudolphi, 1809 
" Qld: THJ, 1912a, p.74 
Contracaecum fricuspe (Gedoelst, 1916) 
Qld: THJ & PMM, 1941c, p.114 


REC. S. AUST. MUS. 19(15): 219-325 

AttgUSQ 1986 

syn. Ascaris sp. Krefft, 1873, p.213 
Qld: Krefft, 1873, p.213 
Synhimantus sp. 
Syn. Dispharynx sp. 

Qld: THJ & PMM, 1941e, p.257 
Microfilaria sp. 

Qld: THJ, 1912a, p.78 


43. Leucocarbo fuseescens (Vieillot) 

Black-Faced Shag (endemic) 
8/8 : T4 C4 N8 A3 

T. Echinoparyphium phalacrocoracis Yamaguti, 
Paryphostomum radiatum (Dujardin, 1845) 
syn. P. tenuicolle (S.J. Johnston, 1917) 
SA: THJ & I. MA, 1942b, p.120 
Peiasiger exaeretus Dietz, 1909 

SA: discarded 
Renicola sp. 

Galactosomum sinuilacte Pearson, 1973 

SA: Pearson, 1973, p.415 
Galactosomum sp. 

SA: AHC (immature) 
Nap/orchis vanissimus Africa, 1938 

Stictodora dipiacantha T.H. Johnston, 1942a 

Hysteromorpha triloba (Rudolphi, 1819) 
SA: THJ, 1942a, p.238 
C. Paradilepis scolecina (Rudolphi, 1819) 
Tas: Prudhoe, 1969, p. 1 90 
Paradilepis sp. 
Qld: C1HUS 

N. Eustrongy tides phalacrocoracis T.H. Johnston & 
Mawson, 194Ie 
SA: THJ & PMM, 1942c, p. 186 
Tas: THJ & PMM, 1945a, p.149 
Capillaria jaenschi T.H. Johnston & Maw.son, 
SA: THJ & PMM, 1945a, p.151 
Capillaria sp. 

Tas: THJ & PMM, 1945a, p.151 
Contracaecum spiculigerum (Rudolphi, 1809) 
SA: THJ & PMM, 1941c, p.lll 
Tas, SA: AHC 
Cosmocephalus jaenschi T.H. Johnston & Mawson, 
Iks: THJ & PMM, 1945a, p.143 
A. Corynosoma clavatum Goss, 1940 

44. Pbaiacroeorax carbo l.inne 
Great Cormorant 
7/7 : T5 C5 N7 

T. Echinoparyphium phalacrocoracis Yamaeuti, 
SA: THJ, 1942a, p.238 
Paryphostomum radiatum (Dujardin, 1845) 
syn. Paryphostomum tenuicolle (S.J. Johnston, 
SA: THJ & LMA, 1942b, p.120 
Petasiger exaeretus Dietz, 1909 
NSW, SA: THJ, 1942a, p.236 
Qld: AHC 
Haplorchis vanissimus Africa, 1938 

SA: AHC (cysts) 
Hysteromorpha triloba (Rudolphi, 1819) 

NSW, SA: THJ, 1942a, p.238 
Schwartzitrema pandubi (Pande, 1939) 
SA: Dubois & LMA, J 972, p.205 
C. Paradilepis scolecina (Rudolphi. 1819) 
SA: Clark, 1957, p.124 


N. Eustrongylides phalacrocoracis T.H. Johnston & 
Mawson, 1941e 
SA: TJH & PMM, 1941e, p.255 
Capillaria jaenschi T.H. Johnston &. Mawson, 
SA: THJ & PMM, 1945b, p.245 
Contracaecum spiculigerum (Rudolphi, 1809) 
Qld, NSW, SA, WA: THJ & PMM, 1941c, p.lll 
Tas: Munday & Green, 1972, p.9 
Tas: AHC 
syn. Ascaris sp. 

NSW: THJ, 1912b, p.108 
Contracaecum sinulabiatum T.H. Johnston & 
Mawson, 1941c 
Qld: THJ & PMM, 1941c, p.113 
Procamallanus murrayensis T.H. Johnston & 
Mawson, 1940b 
SA: THJ & PMM, 1944, p.64 
Cosmocephalus jaenschi T.H. Johnston & Mawson, 
SA: THJ & PMM, 1941e, p.259 
Echinuria squamata (l.instow, 1883) 
SA: THJ & PMM, 1941e, p.257 
Synhimanftts sp. 
Microfilaria sp. 

Qld: Mackerras, 1962, p.436 

45. Phalacrovorax varius (Gmelin) 

Pied Cormorant 

5/5 : T5 CI N4 A3 

T. Renicola sp. 

Galactosomum sinuilacte Pearson, 1973 

Qld, SA: Pearson, 1973, p.415 
Stictodora dipiacantha T.H. Johnston, 1942a 

SA: THJ, 1942a, p.239 




Cardiocephaloides ovicorpus Dubois & Angel, 1972 

SA: Dubois & LMA, 1972, p.204 
Strigeidae, unidentified 
SA: AHC (larval) 
Fellodistomidac, unidentified 
syn. Steringophoridae, unidentified 
* WA: Goss, 1940, p.7 
C. Dilepis maxima Goss. 1940 
WA; Goss, 1940, p.8 
Trypanorhyncha, unidenlified 

SA; AHC (cysts) 
N. Capillaria jaenschi T.H. Johnston & Mawson, 
SA: THJ & PMM, 1945b, p.245 
Contracaecum spiculigerum (Rudolphi, 1S09) 
WA: THJ & PMM, 1941c, p.lll 
A. Corvnosoma clavatum Goss, 1940 
WA: Goss, 1940, p.12 

46. Phalacrocorax sulcirostris (Brandt) 

Little Black Cormorant 

8/8 : T8 C6 N6 A5 

T. Echinochasmus sp. [spines (20)] 
Qld: AHC 
Echinoparvphium phalacrocoracis Yamaguti, 1939a 

SA: THJ, 1942a, p.238 
Parvphostomum radiation (Dujardin, 1845) 
" Qld, Vic, SA, WA; THJ, 1942a, p.236 

Qld: AHC 
syn. P. tenuicotle (S.J. Johnston, 1917) 
* SA: THJ & LMA, 1942b, p.120 
syn. P. phalacrocoracis Goss, 1940 
WA: Goss, 1940, p.2 
Petasiger exaeret us Diet/., 1909 
SA: THJ, 1942a, p.236 
Qld: AHC 
Eehinostomatinae [spines missing) 

Haplorchis sprenti Pearson, 1964 

Qld: Pearson, 1964, p.639 
Haplorchis paravanissimus Pearson & Ow-Yang, 
Qld: Pearson & Ow-Yang, 1982, p.50 
syn.//. vanissimus of Pearson, 1964 (in part) not 
Africa, 1938 
Qld: Pearson, 1964, p.639 
Haplorchis vanissimus Africa, 1938 

Qld: Pearson, 1964, p.633; Pearson & Ow-Yang, 
1982, p.49 
Stictodora diplacanlha T.H, Johnston, 1942a 

Hvstcromorpha triloba (Rudolphi, 1819) 
NSW, SA: THJ, 1942a, p.238 
syn. Diplostomum granulosum Goss, 1940 
WA: Goss, 1940, p.6 
Posthodiplostomum australe Dubois, 1937a 

SA: Dubois & LMA, 1972, p.212 

Diplostomidae, unidentified 

Qld; AHC (immature) 
Schwartzitrema pandubi (Pande, 1939) 

Qld: Dubois & Pearson, 1965, p.89 

SA: Dubois & LMA, 1972, p.205 

C. Paradilepis minima (Goss, 1940) 

SA: Clark, 1957, p.126 
syn. Dilepis minima Goss, 1940 
" WA: Goss, 1940, p.10 

Qld, Vic, SA: AHC 
N. Eustrongylides sp. 

Capillaria jaenschi T.H. Johnston & Mawson, 
1945 b 

SA: THJ & PMM, 1945b, p.245 

Contracaecum spiculigerum (Rudolphi, 1809) 

Qld, SA: THJ & PMM, 194le, p.lll 

syn. Ascaris spiculigerum Rudolphi, 1809 

Qld: THJ, 1912a, p.64 
Sfreptocara recta (Linstow, 1879) 

Desmidocercella sp. 

Microfilaria sp. 

Qld: THJ, 1912a, p.78 
A. Corynosoma clavatum Goss, 1940 


WA: Goss, 1940, p.12 
? Centrorhynchus sp. 


47. Phalacrocorax melanolcucos 


Little Pied Cormorant 

25/25 : T23 C24 N21 A9 

T. Echinoparvphium phalacrocoracis Yamaguti, 
SA: THJ, 1942a, p.238 
Parvphostomum radiatum (Dujardin, 1845) 
' Qld, NSW, Vic, SA; THJ, 1942a, p.233 

syn. Echinochasmus tenuicolits S.J. Johnston, 
NSW: SJJ, 1917, p.206 
Qld: THJ, 1918, p.212 
syn. Parvphostomum tenuicotle (SJJ, 1917) 

SA: THJ & LMA, 1942b, p.120 
syn. Parvphostomum phalacrocoracis Goss, 1940 
WA: Goss, 1940, p.2 
Petasiger exaeret us Diet 2, 1909 
NSW, SA: THJ, 1942a, p.236 
Lehinostomaiidae [spines (27(4))) 


REC S. AUST. MUS. 19()5): 219-325 

August. 1986 

Prosthogonimus vitellatus Nicoll, 1914a 

SA: LMA, 1973, p.859 
Haplorchis pumilio (Looss, 1896) 

Old; Pearson, 1960, p.93; 1964, p.618 
Haplorchis sprenti Pearson 1964 

Old: Pearson, 1964, p.639 
syn. Haplorchis sp. 

Qld: Pearson, I960, p.93 
Procerovum sp. 

Qld: Pearson, I960, p.93 
Stellantchasmus asp'mosus Pearson, 1964 

Qld: Pearson, 1964, p.668 
syn. Stellantchasmus Jalcatus of Pearson, 1960, 
not Onji & Nishio, 1916 

Qld: Pearson, I960, p.93 
Hysteromorpha triloba (Rudolphi, 1819) 

Vic, SA: THJ, 1942a, p.238 
Posihodiptostomum australe Dubois, 1937a 

SA: Dubois & LMA, 1972, p.212 
Schwartzitrema pandubi (Pande, 1939) 

SA: Dubois & LMA, 1972, p.205 
Dolichosaccutus solecarius (S.4. Johnston, 1917) 
syn. Dolichosaccus solecarius S J. Johnston, 1917 

NSW: SJJ t 1917, p.218 

C. Paradiiepis minima (Goss. 1940) 
SA: Clark, 1957, p.I26 
syn. Dilepis minima Goss, 1940 
WA: Goss, 1940, p.10 
Paradiiepis sp. 

SA: Clark, 1957, p.129 
Microsomacanthus cormorant! (Ortlepp, 1938) 
syn. Hymenolepis cormorant! Ortlepp, 1938 
SA: Clark, 1957, p.131 
Woodiandia phalacrocoracis (Woodland, 1929) 
syn. Hymenolepis phalacrocoracis Woodland, 
SA: Clark, 1957, p.132 

Qld, NSW f Vic, SA: AHC 
N. Eustrongylides phalacrocoracis T.H. Johnston & 
Mawson, 1941e 
SA: THJ & PMM, 1941e, p.255 
Capiilaria jaenschi T.H. Johnston & Mawson, 
SA: THJ & PMM, 1945b, p.245 
Contracaecum spiculigerum (Rudolphi, 1909) 

Qld, NSW, SA: THJ & PMM, 1941c, p.lll 
Contracaecum sinulabiatum T.H. Johnston 8c 
Mawson, 1941c 
SA: THJ & PMM, 194k, p.113 
Tetrameres sp. 
Svnhimantus sp. 

SA: THJ & PMM, 1942e, p.71 
Chevreuxia sp. 
Streptocara recta (Linstow, 1879) 

SA: THJ & PMM, 1942e, p.71 
Desmidocercella sp. 

Microfilaria sp. 

NSW: Cleland & THJ, 1912, p.430 

A. Corynosoma clavatum Goss, 1940 
WA: Goss, 1940. p.12 
Qld: AHC 

48. Phaiacroconn sp. 

T. Renicola sp. 

Vic: UMVS 
Stictodora diplacantha T.H. Johnston, 1942a 

Posthodiplostomum australe Dubois, 1937a 


49. Fregata minor (Grnelin) 
Great Frigatebird 

1/1 : N 

C. Tetrabothrius sp. 

Qld: THJ, 1912b, p.108 
Qld: AHC 
N. Contracaecum sp. 
Qld, Vic: AHC 
Seuratia shipleyi (Stossich, 1900) 
Qld: AHC 

50. Fregata artel (Gray) 

Least Frigatebird 

l/l : C N 

C. Tetrabothrius polyorchis Nvbelin, 1917 

WA: Nybelin, 1917, p.15 
N. Seuratia shipleyi (Stossich, 1900) 

Qld: not kept 


51. Phaethon rubricauda Boddaert 

Red-tailed Tropiebird 

1/1 : N 

N. Contracaecum sp. 

SA: AHC (immature) 


52. Ardea pacifiea Latham 
Pacific Heron 

T. Distoma sp. 

NSW: Krefft, 1873, p.213 
C. No record. Dendrouterina australiensis Baer 8c 
Bona, I960, p.13, was attributed to A. pacifiea in 
mistaken identification of 'Grey Heron' 
N. Contracaecum spiculigerum (Rudolphi, 1809) 
Qld: THJ & PMM, 1941c, p.lll 
Porrocaecum reticulatum (Linstow, 1899) 

Qld: THJ & PMM, 1941c, p.115 
Desmidocercella sp. 



Qld: AHC 

'Filarial worms* 

Qld: Maekerras, 1962, p.435 

53. Ardea novaehollandiae Latham 
White-faced Heron 
14/14 : TT2 C12 N12 

T. Eucotvle sp, 

fas: AHC 
Proucetabulorchis dogieti Belopolskaya & 
Bychovskaya-Pavlovskaya, 1954 
Qld: LMA & Pearson, 1977, p.130 
Maritrema oocyst um (Labour, 1907) 

Tas: S.J. Smith, 1981, p.393 
Haplorchis sprenti Pearson, 1964 

Qid: Pearson, 1964, p.639 
Procerovum varium Onji & Nishio, 1916 

Qld: Pearson, 1964, p.654 
Diplostomum galaxiae Smith & Hickman, 1983a 

Tas: Smith & Hickman, 1983a, p.29 
Posthodiplostomum australe Dubois, 1937a 
Qld: Dubois & Pearson, 1967, p.201 
SA: Dubois & LMA, 1972, p.212 
Apharvngostrigea simplex (S.J. Johnston, 1904) 
SA: Dubois & LMA, 1972, p. 197 
Vic, SA: AHC 
syn. Holostomum simplex S.J. Johnston, 1904 
NSW: SJJ, 1904, p.112 
Parastrigea re pens (Chase, 1921) 
syn. Holostomum repens Chase, 1921 
NSW: Chase, 1921, p.500 
Schwartzitrema pandubi (Pande, 1939) 

SA: AHC (cysts) 

SA: AHC (cysts, metacercariae) 
C. Dendrouterinu australiensis Baer & Bona, 1960 
NSW, SA: Baer & Bona, 1960, p. 13; Bona, 
1975, p.91 
Paraditepis urceina Bona, 1975 

SA: Bona, 1975, p.503. 'Grey Heron', quoted 
by Baer & Bona, is Ardea novaehollandiae 
(colloquial use) 
Parvitaenia ardeae (T,H. Johnston, 1913) 
svn. Bancroftiella ardeae T.H. Johnston, 1913 
' Qld: THJ. 1913, p.85; 1912b, p. 107 
Parvitaenia clavipera Baer & Bona, 1960 

Qld: Baer & Bona, 1960, p.13; Bona, 1975, 
syn. Bancroftiella glandularis of THJ, 1912b, (in 
part) not (Fuhrmann, 1905) 
Qld, NSW: THJ, 1912b, p.107 
Parvitaenia paraevctorchida Baer &. Bona, I960 
NSW: Baer & Bona, 1960, p.3; Bona, 1975, 
syn. Bancroj fie/la glandularis of THJ, 1912b, (in 
part), not (Fuhrmann, 1905) 
Qld, NSW: THJ, 1912b, p.107 

NSW, Tas, SA: AHC 
N. Contracaecum spiculigerum (Rudolphi, 1809) 
NSW: THJ & PMM, 1941c, p.Ul 
Contracaecum sp. 

SA: THJ & PMM, 1941c, p.115 

Vic: AHC 
Porrocaecum sp. 

Qld: AHC 
Tetrameres sp. 

Qld, SA: AHC 

54. Ardeola ibis (Linne) 

Cattle Egret 

N. Microtetrameres egretes Rasheed, I960 

55. Egretta alba (Linne) 

Great Egret 
4/3 : T2 CI N3 

T. Apharvngostrigea sp. 

Qld: C1HUS 
Echinoparyphium oxyurum S.J. Johnston, 1917 

Qld: SJJ, 1917, p.201 
Echinostoma sp. 

Qld: THJ, 1912b, p.107 

Patagifer fraternus SJ. Johnston, 1917 

Qld: SJJ, 1917, p.213 
Renicola sp. 

SA: AHC (cysts, metacercariae) 
Haptorchis sprenti Pearson, 1964 

Qld: Pearson, 1964, p.639 
Procerovum varium Onji & Nishio, 1916 

Qld: Pearson, 1964, p.653 
Hetcrophyidae, unidentified 

Posthodiplostomum australe Dubois, 1937a 

SA: Dubois & LMA, 1972, p.212 
Distoma sp. 

NSW: Krefft, 1873, p.213 
C. Anomotaenia asvmmetrica T.H. Johnston, 1913 

Qld: THJ, 1913, p.81 
Parvitaenia glandularis (Fuhrmann, 1905) 
syn. Bancroftiella glandularis (Fuhrmann, 1905) 

Qld: THJ, 1913, p.84 

? Qld: CIHUS 
N. Contracaecum sp. 

SA: THJ & PMM, 1941c, p.114 

Qld: CIHUS (immature) 
Porrocaecum reticulatum (Linstow, 1899) 

Qld: THJ & PMM, 1941c, p.115 

Thetazia sp. 

A. Unidentified 

? loc: CIHUS 

56, Egretta garzvtta (Linne) 
Little Egret 

T Nephrostomum sp. 
Tas: AHC 
Fehinostomatidae [spines (49(4))] 
Tas: AHC 


feEC. S. AUST, MUS. 19(15); 219-325 

AligU&h 19H6 

Diplostomum amygdalum Dubois & Pearson, 1965 
Qld: Dubois & Pearson, 1965, p.90 

Apharyngostrigea simplex (S.J. Johnston, 1904) 
Qld: Dubois & Pearson, 1965, p.79 

57. Egret ta intermedia (Wagler) 
Intermediate Egret 

T, Hu/tlorchis pumilio (Looss, 1896) 

Old: Pearson, 1964, p.618 
Haplorchis sprenti Pearson, 1964 

Qld: Pearson, 1964, p.639 
Procerovum varium Onji & Nishio, 1916 

Qld: Pearson, 1964, p.655 
Diplostomum amygdalum Dubois & Pearson, 1965 

Qld: Dubois & Pearson, 1965, p.90 
Apharyngostrigea simplex (S.J. Johnston, 1904) 

Qld: Dubois & Pearson, 1965, p.79 
C. None recorded. Baer & Bona (1960) and Bona 
(1975) mistakenly quote this bird as host of 
Anomotaenia (/symmetrica THJ, 1913, and of 
Bancroft iel/a glandularis (Fuhrmann) of THJ, 1913, 
but THJ gives host of both species as Herodias 
timorensis Lesson, now Egretta alba. 

SA: Dubois & LMA, 1972, p,206 

Posthodiplostomum australe Dubois, 1937a 

NT: Dubois & LMA, 1972, p.212 
C. Parvitaenia ardeae (T.H. Johnston, 1913) 

SA: Bona, 1975, p.232 

syn. Baneroftiella ardeae T.H. Johnston, 1913 

Qld: THJ, 1913, p.85 
Hvmenolepis sp. 

NSW: T.H. Johnston, 1912b, p. 107 

Qld, NSW: AHC 

Qld, NSW, SA, NT: AHC 
N. Eustrongylides sp. 

Conlraeaecum microcephalism (Rudolphi, 1809) 

Conlraeaecum nycticoracis T.H. Johnston & 
Mawson, 1941c 

NSW: THJ 6c PMM, 1941c, p.lll 
Conlraeaecum \piculigerum (Rudolphi, 1809) 

Qld, SA, NT: AHC 
Desmidocercella sp. 


58. Egretta sacra (Gmelin) 
Eastern Reef Egret 

T. Ausirobilharzia terrigalensis S.J. Johnston, 1917 

Qld: Rohde, 1977, p.39 
C. Baerbonia parviiaeniunca (Baer & Bona, I960) 
Qld: Bona, 1975, p.166 
syn. Valipora parviiaeniunca Baer & Bona, 1960 
' Qld: Baer & Bona, 1960, p.9 

59. Butorides striates (Linne) 
Striated Heron 

4/4 : N 

N. Contracaecum sp. 

60, Mycticomx caledonieus (Gmelin) 

Rufous Night Heron 
3/3 : T2 C2 N3 

T. Haplorchis paravanissimus Pearson & Ow-Yang, 
Qld: Pearson & Ow-Yang, 1982, p.50 
syn. /Y. vanissimus of Pearson, 1964 (in part) not 
Africa, 1938 
Qld: Pearson, 1964, p.633 
Haplorchis vanissimus Africa, 1938 

Qld: Pearson, 1964, p.633 
Procerovum varium Onji & Nishio, 1916 

Qld: Pearson, 1964, p,654 
Stellantchasmus falcatus Onji & Nishio, 1916 

Qld: Pearson, 1964, p.664 
Clinostomum complanatum (Rudolphi, 1814) 
syn. Clinostomum hornum Nieoll, 1914b 
" Qld: Nieoll, 1914b, p.123 
Diplostomum amygdalum Dubois & Pearson, 1965 
Qld: Dubois & Pearson, 1965, p.90 

61. ixobrychus minutus (Linne) 
Little Bittern 

2/1 : N 

N. Contracaecum sp. 

SA: AHC (immature) 
Desmidocercella sp. 
SA: specimen lost 

62. Botaurus poidluptilus (Wagler) 
Australian Bittern 

1/1 ;TCN 

T. Clinostomum complanatum (Rudolphi, 1814) 
syn. Clinostomum hornum Nieoll, 1914b 

Qld: Nieoll, 1914b, p.123 (immature) 
Posthodiplostomum australe Dubois, 1937a 

Qld: Dubois, 1937b, p.34J 
Posthodiplostomum ob/ongum Dubois, 1937a 

Qld: Dubois, 1938, p.291 
Schwartz! frema pandubi (Pande, 1939) 

SA: Dubois & LMA, 1972, p.205 
Hexangium sp. 

Qld: CIHUS (? in food) 
Gyliauchen sp. 

Qld: CIHUS (? in food) 


C. Baneroftiella sp. 

Qld: BM(NH) 

Qld, NSW: AHC 
SA: AHC (eystieereoids) 
N. Capillar ia sp. 
Qld: AHC 
Contracaecum spiculigerum (Rudolphi, 1809) 
SA: THJ & PMM, 194k-, p.lll 



Contracaecum sp., ? microcephalum (Rudolphi, 
A. Arhvihmorhynchus brevis Van Cleave, 1916 


63. Xenorhynchus asiaticus (Latham) 

Black-necked Stork 

1/1 : T C N 

T. Chuunoeephalus ferox (Rudolphi, 1795) 

Qld: Nicoll, 1914b, p.117 
C. Clelandia parva T.H. Johnston, 1909b 

NSW: THJ, 1909b, p.146 
N. Contracaecum sp. 

SA: THJ & PMM, 1941c, p.114 

SA (cage): AHC (larva, encysted) 


64. Plegadis faleinellus (Linne) 

Glossy Ibis 


T. Patagifer bilobus (Rudolphi, 1819) 

Qld: Nicoll, 1914a, p.338 
N. Contracaecum spicuiigerum (Rudolphi, 1809) 
Qld: AHC 
Phvsaiopfera sp. 
* Qld: AHC 

65. Threskiornis acthiopica (Latham) 

Sacred Ibis 

7/7 : T5 C4 N5 A4 

T. Echinostoma acuticauda Nicoll, 1914b 
Qld: AHC 
Patagifer acuminatus S.J. Johnston, 1917 
Qld: SJJ, 1917, p.210 
syn. Echinostoma sp. 

Qld: THJ, 1912b, p-107 
syn. Patagifer bilobus of THJ, 1916, not 
' (Rudolphi, 1819) 
Qld: THJ, 1916, p.47 
Patagifer sp. [spines 2x (30(4))] 

Patagifer sp. [spines 2x (29(3))] 

Vic: AHC 
Platynotrema bitiosum Nicoll, 1914b 

Qld: Nicoll, 1914b, p.U8 
Hvsteromorpha ptataieae Dubinina & Dubinin, 
Qld: Dubois & LMA, 1972, p.208 
Parasthgea sp. 

Qld: Dubois & LMA, 1972, p.198 
Strigea bav/isi Dubois, 1937a 

SA: Dubois & LMA, 1972, p.198 

SA: AHC (cysts) 

C. Paradilepis urceina Bona, 1975 
SA: Bona, 1975, p.503 
Hymenolepididae, unidentified 


Qld, Vic, SA: AHC 
N. Capitlaria sp. 
'Nematoda (? hookworm)' 

Skrjabinociava sp. cf. S. alii Ali, 1968 

Svncuaria contorta (Molin, 1858) 
Vic, SA: PMM, 1982, p.19 
A. Polvmorphus biziuraeTM. Johnston & Edmonds, 
Tks, SA: AHC 

66. Threskiornis spinicollis (Jameson) 
Straw-necked Ibis 

2/1 : N 

T. Echinostoma acuticauda Nicoll, 1914b 

Qld: Nicoll, 1914b, p.110 
Patagifer bilobus (Rudolphi, 1819) 

Qld: Nicoll, 1914b, p.115 

Strigea baylisi Dubois, 1937a 

Qld: Dubois, 1937b, p.235 
Opisthorchis obsequens Nicoll, 1914b 

C. Hvmenolepis ibidis T.H. Johnston, 1913 

Hymenolepis sp. 

Aust: BM(NH) 

Qld; AHC 

Vic: UMVS 
N. Capillaria sp. 

Physaioptera sp. 

Qld: THJ & PMM, 1941e, p.257 

67. Platalea regia Gould 

Royal Spoonbill 

1/1 : A 

T. Orchipedum sufflavum Nicoll, 1914b 

Qld: Nicoll, 1914b, p.108 
Patagifer bilobus (Rudolphi, 1819) 

Qld: SJJ, 1913, p.37] 

? loc: CIHUS 
C. Cyclorchida omalancristrota (Wedl, 1856) 

Qld: THJ, 1913, p.86; Bona, 1975, p.412 

A. Unidentified 

SA: specimens lost. 

68. Platalea flaxipes Gould 

Yellow-billed Spoonbill 

5/5 : T5 C4 N2 A2 

T. Patagifer sp. [spines 2x ? (29(4))| 


REC. S. AUST. MUS. 19(15): 219-325 

August, 1986 

Patagifer sp. [spines 2x (21(1 3))] 

Qld: AHC 
Schwartzitrema pandubi (Pande, 1939) 

SA: Dubois & LMA, 1972, p.205 (cysts) 
Strigea bay list Dubois, 1937a 

SA: Dubois & LMA, 1972, p.198 
C. Paradilepis patriciae Baer & Bona, 1960 

SA: Baer & Bona, 1960, p.17; Bona, 1975, 
Hymenolepis ibidis T.H. Johnston, 1913 

Qld: THJ, 1913, p.88 
Hymenolepis sp. 


Qld, SA: AHC 
N. Contracaecum sp. 


A. Polymorphus biziurae T.H. Johnston & Edmonds, 


69. Anseranas semipalmata (Latham) 

Magpie Goose 

1/1 : T C N 

T. Schistosomatidae (eggs, from nasal mucosa) 

Qld: Blair & Otteson, 1979, p.984 
Notocotylus attenuatus (Rudolphi, 1809) 

Qld: Nicoll, 1914b, p.125 

Typhlocoelum reticulare S.J. Johnston, 1913 

Qld: SJJ, 1913, p. 372 

Echinostoma revolutum (Froelich, 1802) 

Qld: Nicoll, 1914b, p.115 

Qld: AHC 

C. Angularella australis (Maplestone, 1921b)* 
Aust: BM(NH) 
Biuterina sp. 

Aust: BM(NH) 
Cloacotaenia megalops (Nitzsch in Creplin, 1829) 
syn. Hymenolepis megalops (Nitzsch, 1829) 
Qld: THJ, 1913, p.89 
Passehlepis stylosa (Rudolphi, 1810) 

Aust: BM(NH) 
Sobolevicanthus terraereginae (T.H. Johnston, 
syn. Hymenolepis terraereginae T.H. Johnston, 
Qld: THJ, 1913, p.89 

Qld, Vic, NT: AHC 
N. Amidostomum anseris (Zeder, 1800) 
Vic (cage): UMVS 
Epomidiostomum sp. 

* Yamaguti, 1959, p.235, notes that this species cannot 
be referred to Angularella (syn. Angularia) and 
probably belongs to some other genus. 

Heterakis sp. 

Vic (cage): UMVS 
Tetrameres anseranas Mawson, 1979 

NT: PMM, 1979, p.178 

Vic (cage): UMVS 
Tetrameres fissispina (Diesing, 1851) 

Philadelphia Zoo: Canavan, 1931, p.221 
Streptocara crassicauda (Creplin, 1829) 

Vic (cage): UMVS 

70. Dendrocygna arcuata (Horsfield) 
Wandering Whistling-Duck 

T. Schistosomatidae, unidentified 

Qld: Blair & Otteson, 1979, p.984 (eggs from 
nasal mucosa) 
Notocotylidae, unidentified 

Qld: AHC 
Echinostomatinae [spines ? (35(5))] 
Qld: AHC 

C. Ophiotaenia hylae T.H. Johnston, 1912e 

Qld: Maplestone, 1921a, p.404 (? with food) 
Cloacotaenia megalops (Nitzsch in Creplin, 1829) 

Qld: C1HUS 
Diorch is fla vescens ( K re f f t , 1873) 

Qld: Maplestone, 1921a, p.403 
Diplogynia oligorchis (Maplestone, 1922a) 
syn. Coturnia oligorchis Maplestone, 1922a 

Qld: Maplestone, 1922a, p.55 
syn. Diploposthe laevis of THJ, 1913, not (Bloch, 
Qld: THJ, 1913, p.91 
Hymenolepis ibidis (T.H. Johnston, 1913) 

Qld: C1HUS 
? Passerilepis zosteropis (Fuhrmann, 1918) 

Aust: BM(NH) 
Qld: AHC 

71. Dendrocygna eytoni (Eyton) 
Plumed Whistling-Duck (endemic) 

T. Schistosomatidae, unidentified 

Qld: Blair & Otteson, 1979, p.984 (eggs from 
nasal mucosa) 

72. Cygnus atratus (Latham) 

Black Swan (endemic) 

19/19 : T14 C19 N13 A3 

T Trichobilharzia sp. 

SA: THJ, 1941, p.276 
Notocotylus attenuatus (Rudolphi, 1809) 

Qld: Nicoll, 1914b, p.125 
Notocotylus sp. 

Tas, SA: AHC 
Paramonostomum caeci Smith & Hickman, 1983b 

Tas: Smith & Hickman, 1983b, p.86 
Notocotylid, ? sp. B, S.J. Smith, 1981 

Qld, Tas: S.J. Smith, 1981, p.256 
Notocotylidae, unidentified 

NSW, Tas, SA: AHC 
Hyptiasmus magnus S.J. Johnston, 1917 



Vic: SJJ, 1917, p,244 
svn. Monostomum sp. 
* Vic: THJ, 1910, p.97 
Cycioeoelidae, unidentified 

NSW, Vic, SA; AHC 
Echinoparvphium etlisi T.H. Johnston & Simpson, 
SA; THJ & IMA, 1949, p.251 
syn. Echinoparvphium sp. 

? Qld: Verma, 1936, p.155; see THJ & LMA, 
1949, p.253 
Echinoparvphium gizzardai Verma, 1936 

?Qld:* Verma, 1936, p.155; see THJ & LMA, 
1949, p,253 
? Echinosioma minimum Vcrma, 1936 
see Verma, 1936, p. 1 50. We consider the host 
given by Verma as 'Black Swan 1 from Patna, 
Behai, to be an error for Mute Swan, cited as Ihc 
host o\' Echinostoma tongicirrum from the same 
locality (p.152) 
Echinostoma revolution (Froeiich, 1802) 
Qld: Nicoll, 1914b, p.H5 
SA: THJ Sl LMA, 1941c, p.321 
Echinostoma sp. [spines (37(4-5))] 

Tas: AHC 
i'ermatrenta longitestis (Vcrma, 1936) 

Calcutta (? cage): Srivastava, 1972, p.177 
syn. Euparvphium longitestis Vcrma, 1936 
" Calcutta (? Qld): Verma, 1936, p.156; see THJ 
Si LMA, 1949, p.253 
Echinostomatidae, unidentified 

Echinostomatinae [spines 41(5))] 

SA: AHC (immature) 
Psitochasmus oxyurus (Creplin, 1825) 

Tas: 5.1 Smith, 1981, p.181 
Psilostomum sp. A, S.J. Smith, 1981 

las: S .!. Smith, 1981, p.201 (? this host) 
Psilostomum sp. B, S.J. Smith, 1981 

las: S,J. Smith, 1981, p.214 
Psilostomum sp. 

Cvlindrotrema cvgni Angel, 1973 

SA: LMA, 1973, p,854 
Aputemon intermedins (S.J. Johnston, 1904) 
SA: THJ & LMA, 1951, p,67 
svn, Hemistomum intermedium S.J. Johnston, 
NSW: SJJ, 1904, p.110 
Cot vlurus magniacetabulus Dubois & Angel, 1972 
SA: Dubois & IMA, 1972, p.204 



C Armadoskrjabinia globosa (S/potanska, 193!) 
syn. Hymenolepis glohosa Szpotanska, 1931 
Aust: Szpotanska, I93J, p.260 
Qld: BM(NH) 
Austnilin/epts southweili (Szpotanska, 1931) 
syn. Hymenolepis southweili Szpotanska, 1931 

Aust: Szpotanska, 1931, p.260 
syn. Echinorhvnchotueniu nana Maplestone &. 
Southwell, |922b 
Qld: Mapiesmne & Southwell, 1922b, p.193 

Cloucotaenia megalops (Njt/sch in Creplin, 1829) 
Warsaw (eage): Koteehi, 1970, p.332 
Vic, SA: AHC 
Diorchis spiralis Szpotanska, 1931 
Aust: Szpotanska, 1931, p.258 
Diorchis stefanski Czaplinski, 1955 

Warsaw (eage): Koteehi, 1970, p.332 
Drepanidotaenia lanceolatu (Bloch, 1872) 
Aust: Szpotanska, 1931, p.254 
Qld: BM(NH) 
Vie, SA: AHC 
syn. Hymenolepis lanceolata (Bloch, 1782) 
Qld:' Maplestone & Southwell, 1922b, p. 1 97 
Drepanidotaenia rapida (Szpotanska, 1931) 
syn. Hymenotepis rapida Szpotanska, 1931 
Aust: Szpotanska, 1931, p.251 
Drepanidotaenia sp. 

Dicranotaenia coronuta (Dujardm, 1845) 
Warsaw (cage): Koteehi, 1970, p.332 
Hvmenolepis chenopis Palmer, 1981 

Vic: Palmer, 1981, p. 126 
Hymenolepis sp. 

Bengal (cage); Southwell, 1916, p. 11 
Qld: BM(NH) 
Hvmenolepis (sJ.) liophallos Krabbe, 1869 

Europe (cage): Krabbe, 1869, p.291 
"Hymenolepis variabilis n.sp. Maplestone 1 * 

Qld: CIHUS. We have found no record of 
publication of this species by Maplestone. 
Hymenolepis (sJj micrancristrota (Wedl, 1855) 
syn. Taenia micruncristrota Wedl, 1855 
Hungary (cage): Wedl, 1855, p.6 
Monosaccunthes curiosa (Szpotanska, 1931) 
syn. Hvmenolepis curiosa Szpotanska, 1931 
' Aust: Szpotanska, 1931, p.252 
Qld: BM(NH) 
Monosaccunthes kazachstanica (Maksinova, 1963) 

Vie: Palmer, 1981, p. 135 
Monosaccunthes sp. 

Parabisaccanthes bisacculina (Szpotanska, 1931) 
Vic: Palmer, 1981, p.130 
Qld: BM(NH) 
syn, Drepanidotaenia bisacculina Szpotanska, 
Aust: Szpotanska, 1931, p.247 
Tscherlkovilepis krabbei (Kowolewski, 1895) 

Warsaw (cage): Koteehi, 1970, p,332 
Gastrotaenta sp. 

Nematoparataenia puradoxa Maplestone Si 
Southwell, 1922b 
Qld: Maplestone & Southwell, 1922b, p. 189 
Vic, SA: AHC 
Aust: BM(NH) 
CvelophvUidca, unidentified 



REC S. AUST. MUS. 19(15); 219-325 

August. 1986 

NSW: Cleland, 1922, p. 105 

Aust: AM 

Old, NSW, Tas, SA; AHC 

Old: CIHUS (larvae) 

Vic: UMVS 
N. Capillaria obsignata Madison, 1945 

England (cage): Wakelin, 1963, p.38I 
syn. Capillaria anatis (Schrank, 1790) 

England (cage): Wakelin, 1963, p.38l. 
Capillaria eltisi T.H. Johnston & Mawson, 1945b 

SA: THJ & PMM, 1945b, p.247 
Amidostomum cygni Wehr, 1933 

SA: THJ & PMM, 1947, p.550; PMM, 1980, 

Tas: AHC 
Amidostomum sp. 


Vic: AHC 
Pseudamidostom sp. 

Trichostrongylus tenuis 

England (cage): Wakelin, 1963, p.381. 
Heterakis circumvaltata (Linstow, 1906b) 

Germany: Linstow, 1906b, p.251 
Heterakis vesicutaris (Froelich, 1791) 
syn. Heterakis papulosa (Bloch, 1782) 

Berlin Zoo: Schneider, 1866, p.69 
Tetrameres australis T.H. Johnston & Mawson, 

SA: THJ & PMM, 1941e, p.262 

Echinuria uneinata (Rudolphi, 1819) 

Vic: UMVS 

Tas, SA: AHC 
A. Polymorphus sp. 

Corvnosoma sp, 


73. St'wtonetta naevosa (Gould) 

Freckled Duck (endemic) 

19/11 T4 C7 N10 

T. Notocotvlidac, unidentified 

Echinoparyphium sp. [spines (41(4))] 

Psilochasmus sp. 

Psitostomum sp. 

C. Diorchis sp. 

Cloacotaenia sp. 


N. Amidostomum acutum (Lundahl, 1848) 

SA: PMM, 1980, p.9 
Amidostomum 'sp. 6' 

Tetrameres sp. 


Svnhimanius sp. 

74. Coreopsis no\aehullandiae Latham 
Cape Barren Goose (endemic) 

T. Psilochasmus sp. 
Tas: AHC 

C. Fimbriarioides intermedia (Fuhrmann, 1913) 
syn. Ftmbriaria intermedia Fuhrmann, 1913 
Calcutta (cage): Mcggitt, 1933, p.153 
Sobolevicanthus sp. 
N. Amidostomum anseris (Zeder, 1800) 
Tas: PMM, 19S0, p.9 
Heterakis vesicularis (Froelich, 1791) 
syn. Heterakis caudata (Linstow, 1906) 
Calcutta Zoo: Maplcstone, 1932, p.413 
Heterakis chenonettae T.H. Johnston, 1912a 

Tas: AHC 
Heterakis dispar (Schrank, 1790) 
Bass Str: BM(NH) 

75. Tadorna ladornoides (Jardine & Selby) 

Australian Shclduck (endemic) 

5/5 : T2 C3 N3 

T. Echinoparyphium sp. [spines incomplete] 
Echinostoma revolutum (Froelich, 1802) 
syn. 'techmostome larvae" Bradley, 1927 
NSW: Bradley, 1927, p.675 

C. Staphylepis lumelluta (Woodland, 1930) 

syn. Hymenolepis lamellata Woodland, 1930 

London Zoo: Woodland, 1930, p.226 
Hymenolepis sp. 

Vic: AHC 

NSW r , SA, Vic: AHC 
N. Epomidiostomum sp. 

Tetrameres sp. 

Vic: AHC 
Echinuria uneinata (Rudolphi, 1819) 

Vic: AHC 
Streptocara crassicauda (Creplin, 1829) 

Vic: AHC 

76. Tadorna radjah Garnot 
Radjah Shelduck 

1/1 : N 

N. Amidostomum acutum (Lundahl, 1848) 
NT: PMM, 1980, p.9 

77. Anas supereUiosa Gmelin 
Pacific Black Duck 
19/19 : Til C13 N8 

T. Trichobitharzia australis Blair & Islam, 1983 
Qid: Blair & Islam, 1983, p.89 



inchobilharzia sp. 

Qld: Blair & Otteson, 1979, p.983 
Schistosomatidae (eggs, nasal mucosa) 

Qld, NSW, Vic: Blair & Otteson, 1979, p.984 
Notocotvlus attenuatus (Rudolphi, 1809) 

Qld: Nicoll, 1914b, p.127 
Paramonostomum bursae Smith & Hickman, 

Tas: Smith & Hickman, 1983b, p.95 
Paramonostomum caeci Smith & Hickman, 1983b 

Tas: Smith & Hickman, 1983b, p.86 
Notocotylidac, unidentified 

Qld, SA: AHC 
Tvphlocoelum sp. 

Qld, Vic, SA: AHC 
Echinostoma revalutum (Froelich, 1802) 

Qld: SJJ, 1913, p.370 

NSW, SA: THJ & LMA, 1941c, p.321 

Tas: AHC 
syn. Echinostoma sp. 

Qld, NSW: THJ, 1912b, p.108 
Hvpoderaeum sp. 

Qld, SA: AHC 
Patagifer sp. [spines 2x ? (26(3))] 

Echinostomatinae [spines (37(5))] 

Qld, SA: AHC 
Echinostomatinae [spines (37(5))] 

Echinostomatinae [spines (45(5))] 

Psilochasmus oxvurus (Creplin, 1825) 

Tas: SJ. Smith, 1981, p.181 
Psilostomum sp. 

Qld: AHC 
? Prosihogonimus sp. 

Levinseniella tasmaniae (S.J. Smith, 1974) 
syn. Microphallus tasmaniae S.J. Smith, 1974 

Tas: S.J. Smith, 1981, p.107 
Maritrema eatvertense S.J. Smith, 1974 

Tas: S.J. Smith, 1981, p.64 
Apalemon gracilis (Rudolphi, 1819) 

Tas: Smith & Hickman, 1983a, p,23 
Cotylurini, unidentified 

Qld, SA: AHC 
Diplostomidac, unidentified 

C. Cloacotaenia megafops (N'rt/scb in Creplin, 1829) 

Qld: C1HUS 


syn. Hvmenolepis megalops (Nitzsch in Creplin, 

NSW: THJ, 1912b, p. 108 
syn. Taenia cvlindrica Krefft, 1873 

NSW: Krefft, 1873, p.220 
Diorchis flavescens (Krefft, 1873) 

Qld: THJ, 1912a, p.66 

Qld: C1HUS 

syn. Taenia fiuvescens Krefft, 1873 

EA: Krefft, 1873, p.219 

Drepanidotaenia lanceolata (Bloch, 1782) 

Qld: BM(NH) 
Hvmenolepis sp. 

NSW: THJ, 1912b, p.108 

Qld, SA: AHC 
Microsomacanthus collaris (Batsch, 1786) 
syn. Hvmenolepis collaris (Balsch, 1786) 

NSW: THJ, 1912b, p.108 
syn. Taenia bairdii Krefft, 1873 

EA: Krefft, 1873, p.224 
Eimbriaria fasciolaris (Pallas, 1871) 

NSW: THJ, 1912b, p.108 
syn. Taenia pediformis Krefft, 1X73 

EA: Krefft, 1873, p.222 
Eimbriaria sp. 

Diploposthe laevis (Bloch, 1782) 

NSW: THJ, I912d, p.12 
Gastrotaenia sp. 


NSW: Cleland, 1922, p.105 

Tas: Munday & Green, 1972, p.4 

Qld, NSW, Las, SA, NT: AHC 

Vic: MUVS 
N. Ca pillar ia sp. 

NSW: Cleland, 1922, p.105 (AHC) 
Amidostomum acutum (Lundahl, 1848) 

Tas, SA: Mawson, 1980, p.9 
Amidostomum sp. 

Epomidiostomum sp. 


Tas, SA, NT: AHC 
Porrocaecum crassum (Deslongchamps, 1824) 

Tas: AHC 
Coniracaecum microcephalia (Rudolphi, 1809) 

NSW: THJ & PMM, 1941c, p.114 
Tetrameres fissispina (Diesing, 1861) 

SA: THJ & PMM, 1949, p.67 

Tas, NT: AHC 
Phvsalopiera sp. 

NSW: THJ Sl PMM, 1941e, p.257 
Echinuria sp. 


78. Anas gibbcrifrons S. Mullet 

Grey Teal 

14/11 : T5 C8 N6 

T. Trichobithurzia sp. 

NSW: Bearup, 1957, p.163 (eggs in nasal 

Qld, NSW, Vic: Blair & Otteson,. 1979, p.984 
(eggs in nasal mucosa) 
Notocotylidae, unidentified 

Jvphlocoelum sp. 

Cvelococlidae, unidentified 

Echinostoma sp. [spines (35(4 or 5))] 


REC. S. AUST. MUS. 19(15): 219-325 

August, 1986 


Hvpoderaeum sp. 

Echinostomalidae, unidentified 

Echinostomatinae [spines (45(5))] 


C. Cloacotaenia ? megalops 

Diorchis flavescens (Krefft, 1873) 

Qld: Baylis, 1934b, p. 129 

Hymenolepis rohertsi Baylis, 1934b 

Qld: Baylis, 1934b, p.129 
Haploparaxis veitchi Baylis, 1934b 

Old: Baylis, 1934b, p.129 
Staphylepis iamellaia (Woodland, 1930) 

Gastrotaenia sp. 



N. Strongyioides sp. 

Capillaria sp. 


Amidostomum acutum (Lundahl, 1848) 

SA: PMM, 1980, p.9 
Epornidiostomum uncinatum (Lundahl, 1848) 

SA: THJ & PMM, 1942e, p.73 

Vic: UMVS 
Epornidiostomum sp. 

Tetrameres sp. 

SA: THJ & PMM, 1942e, p.71 

Echinuria uncinate (Rudolphi, 1819) 

syn. Echinuria querquedulae T.H. Johnston & 
Mawson, 1942e 

SA: THJ & PMM, 1942e, p.71 
Echinuria sp. 

Streptocara sp. 

SA: THJ & PMM, 1942e, p.72 


79. Anas caslanea (Eyton) 

Chestnut Teal (endemic) 

1/1 : T C N 

T. Hypoderaeum conoideum (Bloch, 1782) 
Tas: AHC 
Hvpoderaeum sp. 
Levinseniei/a tasmaniae (S.J. Smith, 1974) 

syn. Microphallus tasmaniae S.J. Smith, 1974 
Tas: S.J. Smith, 1974, p.202 
Maritrema calvertense S.J. Smith, 1974 
Tas: S.J. Smith, 1974, p.20I 

C. Cloacotaenia megalops {N\t/sch in Creplin, 1829) 
syn. Hvmenoiepis megalops (Nitzseh in Creplin, 
NSW: THJ, 1912a, p.66 
Diorchis flavescens (Krefft, 1873) 

NSW: THJ, 1912d, p.15 
Diorchis sp. 

Diploposthe laevis (Bloch, 1782) 

NSW: THJ, 1912d, p.12 
Microsomacanthus coliaris (Batsch, 1786) 
syn. Hvmenoiepis coliaris (Batsch, 1786) 
NSW: THJ, 1912b, p. 108 
Fimbriaria fasciolaris (Pallas, 1871) 
syn. Taenia pediformis Krefft, 1873 
NSW: Krefft," 1873, p.222 
NSW: THJ, 1912a, p.66 
Vic: AHC 
N. Capillaria anatis (Schrank, 1790) 

G. Brit, (cage): Wakelin, 1965, 0.293 
Capillaria sp. 

Tas: AHC 
Epornidiostomum sp. 

Tas: AHC 
Tetrameres fissispina (Diesing, 1861) 

Tas: AHC 
Echinuria uncinate (Rudolphi, 1819) 
Tas: AHC 

80. Anas rhynchotis Latham 

Australian Shoveler 

5/5 : T2 C5 N5 

T. Schistosomatidae, unidentified 

NSW: Blair & Otteson, 1979, p.984 (eggs from 
nasal mucosa) 
Echinoswmu revolutum (Froelich, 1802) 

SA: THJ & LMA, 1941c, p.321 
Eehinostomatidae, unidentified, 

Echinostomatinac [spines (45(5))] 
C. Cloacoteenia ? megalops (Nitzseh in Creplin, 
Diorchis flavescens (Krefft, 1873) 
syn. Taenia flavescens Krefft, 1873 
NSW: Krefft, 1873, p.219 
Gastrotaenia sp. 


N. Capillaria sp. 
Amidostomum acutum (Lundahl, 1848) 

Tetrameres sp. 
Streptocara crassicauda (Creplin, 1829) 

Streptocara sp. 



Acuariidae, unidentified 

81. Malacorhynchus membranaeeus 


Pink-cared Duck (endemic) 

3/2 : T2 C2 N2 

T. Paramonostomum sp. 

Notocolylidae, unidentified 

Vic, SA: AHC 
Cyclocoelum sp. 

Vic: AHC 
Echinostoma sp. [spines incomplete] 

Vic: AHC (immature) 
? Microphaliidae, unidentifiable 


Vic, SA: AHC 
C. Unidentified 

Vic, SA: AHC 
N. Capillar ia sp. 

Acuariidac, unidentified 

SA: AHC (larva) 

82. Ayihya mist nil is (Eyton) 

5/4 : T3 C4 N2 

T. Schistosomatidae (eggs, nasal mucosa) 

Qld, NSW: Blair & Otteson, 1979, p.984 
Echinostomatidae, unidentified 



SA: AHC (eggs) 
C. Cloacofaeniu, sp. 

Diorchis flavescens (Krefft, 1873) 

NSW: THJ, 1912d, p.15 
Diptogynia oligorchis (Maplestone, 1922) 

Old: C1HUS 
Diplogvnia, sp. 

? Sobolevicanthus sp. 

Diploposthe laevis (Bloch, 1782) 

Qld: Maplestone, 1922a, p.60 

NSW: THJ, 1912d, p.4 

Aust: BM(NH) 
syn. Taenia tuberculata Krefft, 1873 
" NSW: Krefft, 1873, p.215 
Fimbriaria sp. 


SA: not kept 
Gastrotaenia, sp. 


Qld, SA, NT: AHC 

Vic: UMVS 

N, Capillaria sp. 

Epomidiostomum sp. 

NSW: WLRR ('sp. 3') 
Streptocara crassicaucla (Creplin, 1829) 

Tas: AHC 
Streptocara sp. 

Tas: Munday & Green, 1972, p,12 

83. Chenonetta juba ta (Latham) 
Maned Duck (endemic) 

T. Schistosomatidae, unidentified 

Qld: Blair & Otteson, 1979, p.984 (eggs from 
nasal mucosa) 
Echinostomatidae, unidentified 
N. Heterakis chenonettae T.H. Johnston, 1912a 

NSW: THJ, 1912a, p.7l; THJ & PMM, 1941c, 
Amidostomum sp. 

Epomidiostomum sp. 

84. Nettapus coromandelianus (Cimelin) 
Cotton Pygmy-Goose 

T. Cvclocoelum sp. 
Qld: AHC 

85. Nettapus pulchellus Gould 
Green Pvgmy-Goose 

2/\ : C N 

T. Notocotylus attenuatus (Rudolphi, 1809) 

Qld: Nicoll, 1914b, p.125 
Echinostoma revolution (Froelich, 1802) 

Qld: Nicoll, 1914b, p.115 
C. Unidentified 

N. Echinuria uncinata (Rudolphi, 1819) 


86. Oxyum australis Gould 

Blue-billed Duck (endemic) 

5/5 : T4 C5 N2 Al 

T Notocotylidae, unidentified 

Tas, SA: AHC 
Echinostomatinae [spines ? (37(5))] 

SA: AHC (immature) 
Echinostomatinae [spines ? (40(5))] 

Apatemon intermedins (S.J. Johnston, 1904) 

SA: Dubois & LMA, 1972, p.201 

C. Unidentified 

Tas: Munday & Green, 1972, p.4 

Tas, SA: AHC 
N. Capillaria sp. 

Epomidiostomum sp. 

Tas: AHC 


Streptocara crassicauda (Creplin, 1829) 
Tas: AHC 
A. Unidentified 

Tas: specimens lost 

87. Buiuru lobata (Shaw) 
Musk Duck (endemic) 
17/17 : TI6 CI3 N7 A9 

REC. S, AUST. MUS. 19(15): 219-325 August, 1986 



T. Notocotylidae, unidentified 

Echinoparyphium eltisi (T.H. Johnston & Simpson, 

1944) TJ 

SA: THJ & LMA. 1949, p.248 
Echinostoma revolutum (Froelich, 1802) 

Qld, SA: THJ & LMA, 1941c, p.321 A 

Echinostomatinae [spines (35(5))] 

Qld, SA: AHC 
Echinostomatidae [spines (37(5))] 

Echinostomatidae, unidentified 

Psilochasmus sp. 

Apatemon vitelliresiduus Dubois & Angel, 1972 

Qld, SA: Dubois & LMA, 1972, p. 199 


C. Taenia moschata Krefft, 1873 

NSW: Krefft, 1873, p.223 N 

(THJ, 1912d, p.3 noted that specimens lost) 
Cyclophvilidea, unidentified 

CloQcotaenia ? mega/ops (Nitzsch in Creplin, 1829) 

Sobolevicanthus sp. 


Gastrotaenia sp. 


Qld, NSW, Tas, SA: AHC 
N. Capillaria sp. 

Amidostomum biziurae T.H. Johnston & Mawson, 

SA: THJ & PMM, 1947, p.551; PMM, 1959, 

NSW, SA: PMM, 1980, p.9 
Amidostomum sp. (sp. 5) 

Tetrameres biziurae T.H. Johnston & Mawson, 

SA: THJ & PMM, I941e, p.26I 

Streptocara recta (Linstow, 1879) 

Streptocara formosensis Sugimoto, 1930 


A. Polvmorphus biziurae T.H. Johnston & Edmonds, 
SA: THJ & Edmonds, 1948, p.71 

88. Elanus notalus Gould 

Black-shouldered Kite (endemic) 


. Porroccaeum circinum T.H, Johnston & Mawson, 
Vic: AHC 

89. Aviceda subcristutit Gould 
Pacific Baza 

Neodiplostomum spathula (Creplin, 1829) 
syn. Neodiplostomum australiense Dubois, 1937a 
" Qld: Dubois, 1937b, p.337 
Centrorhynchus asturinus (T.H. Johnston, 1913) 
NSW: THJ, 1918, p.215 
syn. Eehinorhynchus bazae Southwell & Macfic, 
Qld: Southwell & Macfie, 1925, p.177 

Qld: C1HUS 

90. Mihus migrans (Boddaert) 
Black Kite 

3/1 C N 


Procvrnea mansioni (Seurat, 1914) 


91. Haliastur sphenurus (Vieillot) 

Whistling Kite 

9/5 : T3 CI Nl 

Echinostomatinae [spines (45(5))] 

SA: AHC (not mature) 
Haplorchis paravanissimus Pearson & Ow-Yang. 
Qld: Pearson & Ow-Yang, 1982, p.50 
syn. H. vanissimus of Pearson, 1964 (in part) not 
Africa, 1938 
Qld: Pearson, 1964, p.633; Pearson & Ow-Yang, 
1982, p.50 
Haplorchis vanissimus Africa, 1938 

Qld: Pearson, 1964, p.633; Pearson & Ow-Yang, 
1982, p.49 
Haplorchis yokogawai (Katsuta, 1932) 

Qld: Pearson, 1964, p.622 
Procerovum varium Onji & Nishio, 1916 

Qld: Pearson, 1964, p.654 
Stellantchasmus falcatus Onji & Nishio, 1916 

Qld: Pearson, 1964, p.665 
Mesostephanus haliasturis Tubangui & Masilungan, 
syn. Mesostephanus minor Dubois & Pearson, 
Qld: Dubois & Pearson, 1965, p.97 
? Diplostomum spathaceum (Rudolphi, 1819) 



Neodiplosromum spathuia (Creplin, 1829) 
Qld: Dubois & Pearson, 1967, p.196 
NSW: Dubois, 1938, p.264 
Neodiplostomum subaequipartitum Dubois & 
Pearson, 1967 
Qld: Dubois & Pearson, 1967, p.199 
SA; Dubois & LMA, 1972, p.210 
Strigea glandulosa Dubois, 1937a 
Qld: Dubois, 1937b, p.244 
SA: Dubois & IMA, 1972, p.198 
Allocreadiidae, unidentified 
SA: AHC (mctaeercariae) 
C. Unidentified 

N. Capillaria sp. 
Contracaecum cevlanicum (Linstow, 1904) 

Procyrnea mansion* (Seurat, 1914) 
syn. Cyrnea mansion? (Seurat, 1914) 
* Qld:' PMM, I968d, p.749 
Microfilaria sp. 

Qld: Mackerras, 1962, p.436 

92. Accipitvr fasciatus (Vigors & Horsfield) 
Brown Goshawk 
10/8 : T2 N8 A3 

T. Apatemon intermedins (S.J. Johnston, 1904) 
SA: Dubois & LMA, 1972, p.201 

C. Unidentified 

Qld, SA: AHC 
N. Microtetrumeres puraccipiter Mawson, 1977 
Qld, Tas, SA, NT: PMM, 1977, p,253 
Microtetrameres sp. 

Vic: AHC 
Spirurida, unidenlified 
NT AHC (larva) 
A. Centrorhvnchus asturinus (T.H. Johnston, 1913) 
Qld: Southwell & Macfie, 1925. p.163 
SA: THJ & Dcland, 1929, p. 148 
syn, Echinorhvnchus sp. 
' NSW: THJ* 1910, p. 100 

93. Accipiler cirrhoccphalus (Vieilloi) 

Collared Sparrowhawk 

5/5 : T2 CI N5 

T. Neodiplostomum spathuia (Creplin, 1829) 
Qld: Dubois & I. MA, 1972, p.209 
Tas: AHC 
syn. Slrigeidae, unidentified 
Tas: Munday & Green, 1972, p.3 
C, Anomotaenia accipiiris T.H. Johnston, 1913 
Qld: THJ, 1913, p.82 

Tas: Munday & Green, 1972, p.4 
N. Physalopiera sp. 

SA: AHC (immature Qs) 
Microtetrameres sp. 

Tas, SA: PMM, 1977, p.257 

Svnhimantus sp. 

Tas: AHC 
Serratospiculum tendo (Nitzsch in Giebel, 1857) 

Qld: Mackerras, 1962, p,432 

? loc: CIHUS 
Serratospiculum guttatum (Schneider, 1866) 


Tas: Munday & Green, 1972, p.12 
Hamatospicutum sp. 

Qld: THJ & PMM, 1941b, p.35 
Microfilaria sp. 

Qld: Cleland & THJ, 1912, p.430 

Qld: Mackerras, 1962, p.436 
Spirurida, unidentified 

Tas: AHC (encysted larvae) 
A. Centrorhvnchus asturinus (T.H. Johnston, 1913) 

Qld: Southwell & Macfie, 1925, p.163 


NSW: THJ & Deland, 1929, p. 148 

94. Accipiter novaehollandiae (Gmelin) 
Grey Goshawk 

N. Porrocaecum circinum T.H. Johnston & Mawson, 
Qld: THJ & PMM, 1941b, p.34 
Thelazia aquilina Baylis, 1934a 
Qld: Bavlis, 1934a, p.150 
Qld: THJ & PMM, 1941b, p.34 
syn. 'nematode* 

Qld: Cleland, 1922, p.107 
Procyrnea mansioni (Seurat, 1914) 
syn. Cyrnea mansioni (Seurat, 1914) 
Qld:' PMM, 1968d, p.749 
Procyrnea ? leptoptera (Rudolph), 1819) 

? Qld: BM(NH) 
Svnhimantus fieldingi (Baylis, 1934a) 
syn, Aeuariu fieldingi Bavlis, 1934a 
" Qld: Baylis, 1934a, p.144 
A. Centrorhvnchus asturinus (T.H. Johnston, 1913) 
Qld:" THJ, 1918, p.215 
Qld: Southwell & Macfie, 1925, p.163 
syn. Gigantorhvnchus asturinus T.H. Johnston, 
Qld: THJ, 1912b, p.108 

95. Haliaeetus leueogaster (Gmelin) 

White-bellied Sea-Eagle 

1/1 : T 

T. Scaphanocephalus australis S.J. Johnston, 1917 
NSW: SJJ, 1917, p.188 
syn. Scaphanocephalus sp. 
? NSW: SJJ, 1914, p.243 
Neodiplostomum spathuia (Creplin, 1829) 
SA: Dubois & LMA, 1972, p.209 
N. Capillaria sp. 
Tas: AHC 
Thelazia aquilina Baylis, 1934a 

Qld: Baylis, 1934a, p.150 
Svnhimantus sp. 
Tas: AHC 


REC. S. AUST. MUS. 19(15): 219-325 

August. 1986 

96. Aquita audax (Latham) 
Wedge-tailed Eagle 

4/1 : N 

T. Neodiplostomum spat hula (Creplin, 1829) 
SA: Dubois, 1982, p. 109 
Strigea glandulosa Dubois, 1937a 
SA: Dubois, 1982, p. 109 
N. Thelazja aquilina Bavlis, 19341 
Qld: Baylis, 1934a, p.150 
Microtetrameres paraccipiter Maw son, 1977 

Tas: AHC 
'Eve worms' 

Qld: Nicoll, 1914c, p.243 

97. Hieraaetus morphnoides (Gould) 

Little Eagle 

N. Porrocciecum circinum T.H. Johnston & Mawson, 
SA: THJ & PMM, 1941b, p.30 

98. Circus assimilis Jardine & Selby 

Spotted Harrier 

1/1 : N A 

C. Cladotaenia feuta Meggitt, 1933 

Calcutta Zoo: Meggitt, 1933, p.161 
Cladotaenia sp. 
Vic: AHC 
N. Porrocaecum circinum T.H. Johnston & Mawson, 
SA: THJ & PMM, 1941b, p.30 
Procyrnea mansioni (Seurat, 1914) 

Microtetrameres circi PMM, 1977 
NT: PMM, 1977, p.255 
A. Unidentified 

NT: specimen not kept 

99. Circus approximate Peales 
Swamp Harrier 

4/4 : T2 C3 N2 

T Opisthorchis sp. 
Neodiplostomum spat hula (Creplin, 1829) 

SA: Dubois & LMA, 1972, p.209 
Parastrigea repens (Chase, 1921) 

SA: Dubois & LMA, 1972, p.198 
Strigea glandulosa Dubois, 1937a 
SA: Dubois & LMA, 1972, p.198 
C. Cladotaenia sp. 
Vic: AHC 

Tas, SA: AHC 
N. Capiltaria sp. 
Porrocaecum sp. 
Tas: AHC 
A. Centrorhynchus asturimts (T.H. Johnston, 1913) 

Qld: THJ, 1918, p.216 


100, Falco subniger Gr&y 

Black Falcon (endemic) 

T. Neodiplostomum spat hula (Creplin, 1829) 

SA: Dubois & LMA, 1972, p.209 
Strigea glandulosa Dubois, 1937a 

SA: Dubois & LMA, 1972, p.199 
C. Cladotaenia sp. 

N. Serratospiculum tendo (Nitzsch in Giebel, 1857) 


101. Falco peregrinus Tunstall 
Peregrine Falcon 
3/3 : Tl CI N3 

T Neodiplostomum spathula (Creplin, 1829) 

SA: Dubois & LMA, 1972, p.209 
Neodiplostomum sp. 

Diplostomidae, unidentified 

Strigeidae, unidentified 

C. Unidentified 

N. Contracaecum sp. 

Microtetrameres raptoris Mawson, 1977 

SA: PMM, 1977, p.255 
Phy salop t era sp. 

Serratospiculum guttatum (Schneider, 1866) 

SA: Bain & PMM, 1981, p.275 
Serratospiculum tendo (Nitzsch in Giebel, 1857) 

SA: Bain & PMM, 1981, p.275 


Vic: UMVS 
syn* S. guttatum (Schneider, 1866) 

SA, NT: THJ & PMM, 1941b, p.34 
Spirurida. unidentified 

SA: AHC (encysted larva) 

102. Falco longipcnnis Swainson 
Australian Hobby 

7/7 : N7 Al 

N. Physaloptera alata Baylis, 1925 

Qld: Baylis, 1925, p.115 
Procvrnea falco (Mawson, 1968d) 

Tas, SA, NT: AHC 
syn. Cyrnea falco Mawson, 1968d 

NT:PMM 1968d, p.752 

Tas: Munday & Green, 1972, p.K) 
Tetrameres sp. 

Microtetrameres raptoris Mawson, 1977 


24 > 

NT: PMM, 1977, p.255 
Cieopetitia falco Mawson, 1966 

NT: PMM, 1966, p.717 
Svnhimantus falco Mawson, 1982 

Serratospiculum guttatum (Schneider, 1866) 

SA, WA: TH.I & PMM, 1941b, p.34 

Tas: Mundav & Green, 1972, p.12 

SA: Bain &PMM, 1981, p.273 

Vic, Tas, NT: AHC 
syn. S. attenuation (Rudolphi, 1819) 

Qld: Baylis, 1925, p.112 
Serratospiculum tendo (Nitzseh in Giebe!, 1857) 

Qld: BM(NH) 

Splendidofilariinae, unidentified 

filarial worms' 

Tas: Cleland, 1922, p.107 
Spirurida, unidentified 

NT: AHC (encysted larvae) 
A, Unidentified 

NT: specimen not kept 

103. Falco hypoleucos Gould 
Grey Falcon (endemic) 

C. Unidentified 

N. Hamaiospiculum sp., near H.quadridens (Molin, 
Serratospiculum guttatum (Schneider, 1866) 

Microfilaria sp. 
Qld: Breinl, 1913a, p.34 (CIHUS) 

104. Falco berigora Vigors & Horsfield 

Brown Falcon 

13/11 : CI Nil A6 

T. Episthmium prosthovitellatum (Nicoll, 1914b) 
syn. Echinochasmus prosthoviiellatus Nicoll, 
Qld: Nicoll, 1914b, p.115 
Psilochasmus sp. 
Opisihorchis obsequens Nicoll, 1914b 

Qld: Nicoll, 1914b, p.106 
Strigea glundulosa Dubois, 1937a 

syn. Strigea falconis Dubois, 1937b (not Szidat, 
Aust. (? Qld): Dubois, 1937b, pp.244,247 
Strigea sp., indeterminable 
C. Unidentified 

Tas, SA, NT: AHC 
Tas: Cleland, 1922, p.] 05 (AHC) 
Tas: Munday & Green, 1972, p.4 
N. Capillaria sp. 

Porrocaecum angusficolle (Molin, 1860) 

Procvrnea falco (Mawson, 1968d) 
Tas, NT: AHC 
syn. Cyrnea falco Mawson, 1968d 
SA: PMM. 1968d, p.752 
Procyrnea mansion! (Seurat, 1914) 
syn. Cvrnea mansion! (Seurat, 1914) 
" NSW: PMM, 1968d, p.749 
Procyrnea ? leptoptera (Rudolphi, 1819) 

Qld: BM(NH) 
Tetrumeres sp. 
Microtetrameres raptoris Mawson, 1977 

SA, NT: PMM, 1977, p.255 
Svnhimantus flinders! (T.H. Johnston & Mawson, 
syn. Acuaria flindersi T.H. Johnston & Mawson, 
Tas: THJ & PMM, 1941b, p.3l 
syn. 'nematode' 
Tas: Cleland, 1922, p.l()5 (AHC) 
Svnhimantus falco Mawson, 1982 
SA: PMM, 1982, p.24 
las: AHC 
Physaloptera hieracidiae T.H. Johnston & Mawson, 
Tas: THJ & PMM, 1941b, p.31 
Bancroftinema dentatum T.H. Johnston & 
Mawson, 1941b 
Qld: THJ & PMM, 1941b, p.33 
Thelazia aquilina Baylis, 1934a 
Qld: Bavlis, 1934a, p.150 
Serratospiculum guttatum (Schneider, 1866) 
syn. Filaha guttata Schneider, 1866 
SA: Schneider, 1866, p.80 
Diplotriaena falconis (Connal, 1912) 

SA: Bain & Mawson, 1981, p.283 
'Eye worms' 

Qld: Nicoll, 1914c, p.244 
A. Centrorhynchus falconis (T.H. Johnston & Best, 
syn. Gordiorhynchus falconis T.H. Johnston & 
Best, 1943 
NT: THJ & Best, 1943, p.229 
Centrorhynchus astuhnus (T.H. Johnston, 1913) 
Qld: Southwell & Macfie, 1925, p. 164 
NSW: THJ & Deland, 1929, p.148 

105. Falco cenchroides Vigors & Horsfield 

Australian Kestrel 

7/5 : CI A4 

C. Unidentified 

N. Procvrnea falco (Mawson, 1968d) 
Procvrnea paraleptoptera (T.H. Johnston & 
Mawson, 1941b) 
syn. Cyrnea paraleptoptera T.H. Johnston & 
Mawson, 1941b 


REC. S. AUST. MUS. 19(15); 219-325 

August, 1986 

Qld: THJ & PMM, 1941b, p.32 
Microtetrameres raptoris Mawson, 1977 

SA: PMM, 1977, p.255 
Synhimantus falco Mawson, 1982 
SA: PMM, 1982, p.24 
A. Centrorhvnchus asturinus (T.H. Johnston, 1913) 
Qld,* SA: AHC 


106. Leipoa oceliata Gould 
Mallee Fowl (endemic) 

2/2 : C2 N2 

C. Raiilieiina leipoae T.H. Johnston & Clark, 1948b 
SA: THJ & Clark, 1948b, p.88 
Raiilieiina sp. 

SA: THJ & Clark, 1948b, p.90 

N. Leipoanema eilisi THJ & PMM, I942e 
SA: THJ & PMM, 1942e, p.73 

107. A tectum tat ha mi Gray 
Australian Brush Turkey (endemic) 

C. Davainea sp. 

Qld: AHC 
Raiilieiina sp. 

SA (cage): AHC 
Qld: AHC 
N. Capillaria sp. 
Qld: AHC 
Svngamus sp. 

Qld: AHC 
Odontoterakis bancrofti (T.H. Johnston, 1912a) 
Qld: AHC 
syn. Heterakis bancrofti T.H. Johnston, 1912a 
" Qld: THJ, 1912a, p.72 
Ascaridia catheturina (T.H. Johnston, 1912a) 
Qld: THJ & PMM, 1942a, p.93 
syn. Heterakis catheturinusTM. Johnston, 1912a 
' Qld: THJ, 1912a, p.73 
Gongvlonema alec turae T.H. Johnston & Mawson, 
Qld: THJ & PMM, 1942a, p.92 
Lissonema sp. 
Qld: AHC 
A. Mediorhynchus aleciurae (T.H. Johnston & 
Edmonds, 1947) 
syn. Empodius aleciurae T.H. Johnston & 
Edmonds, 1947 
Qld: THJ & Edmonds, 1947, p.557 
syn. Echinorhvnchus sp. 
Qld: THJ, 1912b, p.106 


108. Coturnix novaezelandiae Quoy & 


Stubble Quail (endemic) 

5/1 : C 

C. Unidentified 


109. Coturnix ypsitophora Bosc 
Swamp Quail 
10/9 : Tl N8 

T. Echinostomatidae [spines missing] 

Tas: discarded 
C. Metroiiasthes sp. 

Vic: AHC 
N. Tetrameres sp. 

Tas: AHC 
Cyrnea co/ini Cram, 1927 

Tas: AHC 
Cheilospirura gruveli (Gendre, 1913) 

Tas: PMM, 1982, p.23 
A. Unidentified 

SA (cage): AHC (cyst) 


110. Turnix castanota (Gould) 

Chestnut-backed Button-quail (endemic) 

2/2 : Tl A2 

T Skr/ubinosomum mawsoni Angel & Pearson, 1977 
SA: LMA & Pearson, 1977, p.124 

A. Oncicola pomaiostomi (T.H. Johnston & 
Cleland, 1912) 
SA: AHC (cyst) 

111. Turnix velox (Gould) 
Little Button-quail (endemic) 

4/1 : N 

N. Acuariidae. unidentified 

A. Oncicola pomatostomi (T.H. Johnston & Cleland, 
syn. Otigacanfhorhvnchus pomatostomi 
(T.H. Johnston & Cleland, 1912) 
Qld: Hall Exped. Report, 1974, p.35 (cyst) 

112. Turnix pyrrhothorax (Gould) 
Red-chested Button-quail (endemic) 

N. Capillaria sp. 

SA (cage): AHC 



113. Pedionomus torquat us Gould 

Plains Wanderer (endemic) 
1/1 : A 

A. Oncicola pomutostomi (T.H. Johnston & Cleland, 
syn. Ech'tnorhynchus pomutostomi T.H. 
Johnston & Cleland, 1912 
SA: THJ & Deland, 1929, p.148 (cyst) 


114. Rallus phiiippensis Linne 

Buff-banded Rail 

4/1 ; T 

T. Leucochloridium sp. 

A. Unidentified 

Qld: AHC (part of worm only) 

115. Rallus pvctoralis Temminck 
Lewin's Rail 
3/3 : Tl N2 

T. Unidentified 

Tas: specimen not kept 

N. Capilloriu sp. 

Tas: Munday & Green, 1972, p.6 
Tas: AHC 

116. Par ana pusilla (Pallas) 

^aillon's Crake 

3/3 : Tl N2 

T. ? Catatropis sp. 

Echinostoma sp. [spines missing] 

Echinostomatinae [spines (45(4-5))] 

SA: AHC (immature) 
N. Capillar ki sp. 

Tetrameres globosa (Linstow, 1879) 

SA: PMM, 1979, p.180 

117. Porzana fluminca Gould 
Australian Crake (endemic) 

14/13 : T3 N13 

T. ? Catutropis sp. 

Cyclocoelidae, unidentified 

SA: AHC (larvae) 
Echinostomatinae [spines missing] 

SA: discarded (immature) 
N. Capillaria sp. 

Porrocaecum ensicaudatum (Zeder, 1800) 

Tetrameres globosa (Linstow, 1879) 

SA, NT: PMM, 1979, p. 180 
Acuariidae, unidentified 
SA: not kept (larva) 

118. Porzana tabuensis (Gmelin) 

Spotless Crake 

2/2 : N 

N. Tetrameres globosa (Linstow, 1879) 
SA: PMM, 1979, p.180 

119. Gallinula mortierii (Du Bus) 
Tasmanian Native-hen (endemic) 

5/3 : N 

N. Capillaria sp. 
Tas: AHC 
Tetrameres globosa (Linstow, 1879) 
Tas: AHC 
syn. Tetrameres sp. 

Tas: Munday & Green, 1972, p.ll 

120. Gallinula ventralis Gould 
Black-tailed Native-hen (endemic) 

7/4 : T2 N2 

T. ? Catatropis sp. 

? Eehinostomam hilliferum Nicoll, 1914b 

Qld: AHC 
Echinostomatidae, unidentified 

SA: believed lost (immature) 
C. Unidentified 

Qld: AHC 
N. Amidostomum tribonvx Mawson, 1980 

SA: PMM, 1980,' p.10 

121. Gallinula tenebrosa Gould 

Dusky Moorhen 

12/7 : T7 C2 N2 

T Catatropis gallinulae T.H. Johnston, 1928 

SA: THJ, 1928, p.135 
? Paramonostomum sp. 

Echinostoma australe T.H. Johnston, 1928 

SA: THJ, 1928, p.138 
Echinostoma bancrofti T.H. Johnston, 1928 

Qld: THJ, 1928, p.140 
? Echinostoma revolution (Froelich, 1802) 

SA: discarded 
Echinostoma sp. [spines (37(5))] 

Qld, SA: AHC 

SA: AHC (immature) 

C. Unidentified 

Qld, SA: AHC 

N. Capillaria sp. 
Amidostomum sp. 


KLC. S« AUST. MUS. 19(15): 219-325 

August, 1986 

122. Porphyria porphyrin (Linne) 

Purple Swamphen 

18/5 : T4 Nl 

T. Echinostoma hilliferum Nicoll, 1914b 

Qld: Nicoll, 1914b, p.112 

Qld, SA: AHC 
Cvciocoelum sp. 

Echinostoma sp. [spines (35(5))] 

Echinostomatinae [spines 35(5))] 

Echinostomatinae [spines ? (27(4))] 

Distoma sp. 

NSW: Krefft, 1873, p.213 
C. Unidentified 


N. Capillar ia sp. 

? loc: AHC 
Tetrameres giobosa (Linstow, 1879) 

123. Fulka atra Linne 
Eurasian Coot 

3/3 : Tl N2 

T. Echinostoma hilliferum Nicoll, 1914b 

Qld: Nicoll, 1914b, p.112 
'Echinostome larvae* 

NSW: Bradley, 1926, p.575; 1927, p.675 
Psiiochasmus oxyurus (Creplin, 1825) 

Tas: S.J. Smith, 1981, p. 1 8 1 
Atriophallophorus coxieliae S.J. Smith, 1974 

Tas: SJ. Smith, 1981, p.181 
C. Hymenolepis sp. 

Qld: AHC 

NSW: Bradley, 1927, p.675 

Qld, SA: AHC 
N. Amidostomum fulica (Rudolphi, 1819) 

Vic: UMVS 

Pelecitus fuiicaeatrae (Diesing, 1861) 

Vic: UMVS 

NSW: Bradley, 1927, p.675 (non-bursate) 


124. Grus rubicundus (Perry) 

T. Allopyge antigones S.J. Johnston, 1913 
Qld: SJJ, 1913, p.375 
syn. Ynonostome' 

Qld: Nicoll, 1914c, p.244 
Echinostoma austraiasianum Nicoll, 1914a 

Qld: Nicoll, 1914a, p.337 
Prosthogonimus sp. 
Qld: AHC 

N. Porrocaecum serpentulum (Rudolphi, 1809) 
syn. Ascaris serpentina Rudolphi, 1809 
Germany (Mus.): Linstow, 1899, p.7 
Asearidia stroma (Linstow, 1899) 
Qld: UQDP 

125. Grus antigone (Linne) 
Sarus Crane 

N. Asearidia stroma (Linstow, 1899) 
NSW (cage): AHC 


126. Ardeotis australis (Gray) 

Australian Bustard (endemic) 

1/1 : C 

T. Philophthulmus sp. 

C. Jdiogenes sp. 

Vic (cage): AHC 
Ascometra sp. 

Vic (cage): AHC 

Qld, Vic (cage), SA (cage), NT: AHC 
Vic (cage): UMVS 
N. Hartertia sp. 

Exeisa dentifera (T.H. Johnston & Mawson, I941d) 
syn. Cyrnea dentifera T.H. Johnston & Mawson, 
NT: THJ & PMM, l941d, p.255 


127. Irediparra gallinacea (Temminck) 

Comb-crested Jacana 
1/1 : T 

T. ? Echinostomatidae, unidentifiable 
NT: AHC (fragment) 


128. Burhinus magnirostris (Latham) 

Bush Thick-knee 

T Notocotylus attenuatus (Rudolphi, 1809) 

Qld: Nicoll, 1914b, p.125 
Echinostoma ignavum Nicoll, 1914b 

Platynotrema hiliosum Nicoll, 1914b 

Qld: Nicoll, 1914b, p.118 
Platynotrema jecoris Nicoll, 1914b 

Qld: Nicoll, 1914b, p.120 
Strigea nicolii Dubois, 1937a 

Qld: Dubois, 1937b, p.241 
C. Angularella austraiis (Maplcstone, 1921b)* 

*Yamaguti, 1959, p.235, notes that this species cannot 
be referred to Angularella (syn. Angularia) and 
probably belongs to some other genus. 



syn. Angufaria uustralis Maplestone, 1921b 
" Qld: Maplestone, 1921b, p,407 
Choanotaenia sp. 

Aust: BM(NH) 
Pa rictero taenia sp. 

Aust: BM(NH) 
Qld: AHC 


129. Haematopus longirostris Vieillot 
Pied Oystercatcher 

C, Unidentified 

Qld, SA: AHC 
A. Prosthorhvnchus sp. 

Tas: AHC 

130. Haematopus fuliginosus Gould 

Sooty Ovstercateher (endemic) 

1/1 : C N A 

C, Unidentified 

N. Sciadocara sp. 

A. Fi/icollis sphaerocephalus (Bremser, 1819) 

Tas, SA: AHC 


131. Yanellus miles (Boddaert) 

Masked Lapwing 
11/11 : CI N10 A2 

T. Notocotylus ; attemtatus (Rudolphi, 1809) 
Qld: Nicoll, 1914b, p.125 
Cvclocoelum obscurum (Leidy, 1887) 

Qld: Hall Exped. Report, 1974, p.354 
Haematrofrephus consimilis Nicoll, 1914b 

Qld: Nicoll, 1914b, p.124 
Plagiorehiidae, unidentified 

syn. Lepodermalidae, unidentified 
Qld: Nicoll, 1914b, p.106 
Echinostotna ignavutn Nicoll, 1914b 

Qld: Nicoll, 1914b, p.114 
Echinosioma sp. [spines ? (35(5))] 
Tas: AHC 
G. An&ularella sp. 
Choanotaenia southwel/i Fuhrmann, 1932 

syn. Monopylidiutn macracanthum of 
Maplestone & Southwell, 1923, not 
Fuhrmann, 1907 
Qld: Maplestone & Southwell, 1923, p.322 
Lapwim>iu adelaidae Schmidt, 1972 

Qld: Schmidt, 1972, p.1085 
Gyrocoelia australiensis T.H. Johnston, 1910 
Qld: Maplestone & Southwell, 1922a, p.61 
Qld, SA: Schmidt, 1972. p.1089 
syn. Gyrocoelia sp. 

Qld: THJ, 1914a, p. 108 
Gyrocoelia sp. 

Infula hurhini Burt, 1939 

Qld, SA: Schmidt, 1972, p.1038 
syn. 'Shipleva lobivanellus Maplestone' 


? NSW: Bradley, 1927, p.675 

Tas: Mundav & Green, 1972, p.6 

Qld, Tas, SA: AHC 
N. Capillaria triloba Linstow, 1875 

SA: PMM, 1968b, p.279 

Tas: Munday & Green, 1972, p.6 
Porrocaecum lobibicis Mawson, 1968b 

SA: PMM, 1968b, p.284 

Tas: Munday & Green, 1972, p.9 

Tas, Vic: AHC 
Tetranieres lobibicis Mawson, 1968b 

SA: PMM, 1968b, p.300 

Tas: AHC 
Tetrameres sp. 

Tas: Munday & Green, 1972, p.ll 
Sireptocara crassicauda (Creplin, 1829) 

Tas: Munday & Green, 1972, p.12 
Streptocara sp. 

Tas: AHC 
Psettdaproctella ? sp. 

Qld: PMM, 1968b, p.302 
Microfilaria sp. 

Qld: Mackerras, 1962, p.436 
A. Prosfhorfivnchus sp. 

Tas: AHC 

132. \anellus tricolor (Vieillot) 

Banded Lapwing (endemic) 

6/4 : C3 Nl Al 

C. Paricterotaenia zoniferae (T.H. Johnston, 1912c) 
syn. Choanotaenia zoniferae T.H. Johnston, 
NSW: THJ, 1912c, p.212 
N. Unidentified 

NT: specimen lost 

A. Unidentified 

SA: specimen lost 

133. Pluvialis dominica (P.L.S. Miillcr) 
Lesser Golden Plover 

T. Acanthoparvphium spinulosntn S.J. Johnston, 
NSW: SJJ, 1917, p.198 
Tas: AHC 
syn. Echinostotna sp. 
" NSW: THJ. 1910, p.93 
Echinostomatidae, unidentified 

Tas: Munday & Green, 1972, p.3 
Levinsemella howensis S.J. Johnston, 1917 
NSW: SJJ, 1917, p.221 
NSW: Pearson & Deblock, 1979, p.32 


REC. S. AUST. MUSL 19(15): 219-325 

August, 1986 

Levinsenietla microovata Belopolskaya, 1958 
Qld; Deblock & Pearson. 1970. a 777 
Microphallus papi/iornatus Deblock & Pearson, 
Qld; Deblock & Pearson, 1969, p.396 
Microphallus vaginosus Deblock & Pearson, 1969 
Qld: Deblock & Pearson, 1969, p.403 
C Unidentified 


134. Erythrogonys ductus Gould 
Red-kneed Dotterel 
10/10 : T6 CIO Nl 




Notoeotylidac, unidentified 

SA: AHC (immature) 
Echinostomatidae [spines ? (35-37(5))] 

SA: AHC (immature) 
Echinostomatidae [spines ? (33(4))] 

SA: AHC (immature) 

SA: AHC (immature) 

Acuaridae, unidentified 

SA: AHC (larva) 

135. Charadrius rubricollis Gmelin 

Hooded Plover (endemic) 

2/2 : N2 A2 




Athophallophorus coxiellae SJ. Smith, 1974 

Tas: S.J. Smith, 1974, p.200 (immature) 
Levinsenietla (asmaniae (S.J. Smith, 1974) 
syn. Microphallus tasmaniae SJ. Smith, 

Tas: S.J. Smith, 1974, p.203 
Maritrema catvertense S.J. Smith, 1974 

tfcs: S.J. Smith, 1974, p.2()l 
Stettocaronema charadrii Mawson, 1968b 

SA: PMM, 1968b, p.294 
Skrjabinoclava horhcla (Rudolphi, 1809) 

SA: PMM, 1968b, p.292 

A. Plagiorhynchus charadrii (Yamaguti, 1939b) 

syn. Prosthorhynchus charadrii Yamaeuti, 1939b 
SA: THJ & SJE, 1947, p.561 

136. Charadrius mongolus Pallas 
Mongolian Plover (migratory) 

T. Endocofyle incana Belopolskaya, 1952 

Qld: Deblock & Pearson. 1968b, p.463 
Gynaecotv/a bhsbanensis Deblock & Pearson, 
Qld: Deblock & Pearson, 1968a, p.141 
Levinsenietla microovata Belopolskaya, 1958 

Qld: Deblock & Pearson, 1970, p.777 
Levinsenietla monodactyla Deblock & Pearson, 
Qld: Deblock & Pearson, 1970, p.774 
Maritrema eroliae Yamaguti, 1939a 

Qld: Deblock & Pearson, 1968b, p.458 
Microphallus paptllomatus Deblock & Pearson, 

Qld: Deblock & Pearson, 1969, p.396 
Microphallus sp. 

Qld: Deblock tk Pearson, 1969, p.400 
C, Unidentified 

Tas, SA: AHC 

137. Charadrius bicinctus Jardine 8c Selbv 

Double-banded Plover (migratory) 

2/2 : CI N2 

C. Unidentified 

N. Capiitaria sp. 

Streptocara crassicauda (Creplin, 1829) 


138. Charadrius leschenauitii Lesson 
Large Sand Plover (migratory) 

T. Acanthoparvphium marilae Yamaguti, 1934 
Acanthoparvphium spinulosum S.J. Johnston, 1917 

Qld: Bearup, 1960, p.219 
Maritrema sp. 
C. Unidentified 


139. Charadrius ruficapillus Tcmminck 

Red-capped Plover (endemic) 

17/14 : T2 C10 N10 A6 

T. Acanthoparvphium spinulosum S.J. Johnston, 

Athophallophorus coxiellae S.J. Smith, 1974 

Tas: S.J. Smith, 1974, p.200 (immature) 
Levinsenietla tasmaniae (S.J. Smith, 1974) 
syn. Microphallus tasmaniae S.J. Smith, 1974 

Tas: S.J. Smith, 1974, p.203 
Mierophallidae, unidentified 

Tas: Munday & Green, 1972, p.3 

C Unidentified 

Tas: Munday & Green, 1972, p.4 

Tas, SA: AHC 
N. Capiitaria sp. 

Tetrameres nouveii (Sen rat, 1914) 

SA: PMM, 1968b, p.297 
Tetrameres sp. 

Tas: Munday Sc Green, 1972, p.ll 
Echinuha hetewbrachiaia Wehr, 1937 

SA: PMM, 1968b, p.289 
Skrjabinoclava horrida (Rudolphi, 1809) 

SA: PMM, 1968b, p.292 

Tas: AHC 
syn. Skrjabinoclava sp. 

Tas: Munday & Green, 1972, p.12 
Streptocara crassicauda (Creplin, 1829) 

SA; PMM, 1968b, p.293 

Tas: AHC 



syn. Streptocara sp. 
Tas: Munday & Green, 1972, p. 12 
Spirurida, unidentified 
A. Plagiorhynchus charadrii (Yamaguti, 1939b) 
Tas, SA: AHC 

140. Charadrius melanops Vieillot 

Black-fronted Plover (endemic) 

12/7 : T3 C3 Nl 

T. Echinostomatidae [spines (37(5))] 
SA: AHC (immature) 
Echinostomatidae [spines ? (41(5))] 

SA: AHC (immature) 
Levinseniella tasmaniae (S.J. Smith, 1974) 
syn. Microphallus tasmaniae S.J. Smith, 1974 
Tas: S.J. Smith, 1974, p.203 
Maritrema calvertense S.J. Smith, 1974 
Tas: S.J. Smith, 1974, p.202 
C. Unidentified 

N. Chevreuxia australis T.H. Johnston & Mawson, 
SA: PMM, 1968b, p.292 

141. Peltohyas australis Gould 
Inland Dotterel (endemic) 

C. Unidentified 

A. Unidentified 

Aust: AHC (fragments) 


142. Himantopus leucoeephalus (Gould) 

White-headed Stilt 

l/l : C N 

T. Trichobilhurzia sp. 


Aust: Bearup & Langsford, 1966, p.523 
Bilharziellinae, unidentified 

Haematotrephus adelphus S.J. Johnston, 1917 

SA: SJJ, 1917, p.24l 

syn. Monostomum sp. 

SA: THJ, 1910, p.94 

C. Davainea himantopodis T.H. Johnston, 1911 
syn. Davainea sp. 

SA: THJ, 1910, p.95 

Diorehis flavescens (Krefft, 1873) 

Ilvmenolepis sp. 

SA: THJ, 1910, p.95 
Acoleus hedlevi T.H. Johnston, 1910 

SA: THJ,' 1912d 

syn. Taenia rugosa Krefft, 1873 

NSW: Krefft, 1873, p.223 
Gyrocoelia australiensis (T.H. Johnston, 1910) 

NSW: THJ, 1912d, p.28 
svn. Dilepis australiensis T.H. Johnston, 1910 

NSW: THJ, 1910, p.95 
syn. Taenia coronata Krefft, 1873 

NSW: Krefft, 1873, p.220 
Infula burhini Burt, 1939 

'Aust': Voge & Read, 1954, p.483 


N. Capillaha triloba Linstow, 1875 

SA: PMM, 1968b, p.279 
Amidostomum acutum (Lundahl, 1848) 

NT: PMM, 1980, p.9 
syn. A. chevreuxi Seurat, 1918 

SA: PMM, 1968b, p.283 
Contracaecum sp. 

SA: PMM, 1968b, p.287 
Teirameres nouveli (Seurat, 1914) 

SA: PMM, 1968b, p.297 

? Chevreuxia revoluta (Rudolphi, 1819) 


143. Cladorhynchus leucoeephalus 


Banded Stilt (endemic) 

7/7 : T5 C4 N6 

T. Bilharziellinae, unidentified 

Notocotylidae, unidentified 

Haematotrephus adelphus S.J. Johnston, 1917 

?Omphalometridae, unidentified 

Acanthoparyphium sp. [spines (23(0))] 

C. Davainea sp. 

SA (cage): AHC 
Diorehis flavescens (Krefft, 1873) 

Gyrocoelia sp. 


SA: AHC (immature) 
N. Tetrameres cladorhvnchi Mawson, 1968b 

SA: PMM, 1968b, p.299 
Echinuria heterobrachiata Wehr, 1937 

SA: PMM, 1968b, p.289 

144. Recurvirostra novaeholiandiae 


Red-necked Avocet (endemic) 

4/4 : Tl C4 N3 

Notocotylidae, unidentified 


REC, S. AUST, MUS. 19(15): 219-325 

August, 1986 


Cyclocoelidae, unidentified 

Hymenolepis sp. 

Davainia sp. 

Himantocestus sp. 


Capillaria triloba Linstow, 1875 

SA: PMM, 1968b, p.279 
Capillaria recurvirostrae Mawson, 1968b 

SA: PMM, 1968b, p.280 
Tetrameres nouveti (Seurat, 1914) 

SA: PMM, 1968b, p.297 
Chevreuxia revolula (Rudolphi, 1819) 

Echinuria heterobrachiata Wchr, 1937 


148. Tringa hypoleucos Linne 
Common Sandpiper (migratory) 

T. Echinoparyphium sp. 

? loc: CIHUS 
Echinostoma sp. 

? loc: CIHUS 
Echinostomatidae [spines missing] 

SA: discarded (very young) 
Platynotrcma sp. 

Mariirema sp. 

C. Haploparaxis australis (T.H. Johnston, 




149. Tringa nebularia (Gunnerus) 

Greenshank (migratory) 

5/4 : Tl C3 Nl 


145. Numenins madagascariensis (Linne) 


Eastern Curlew (migratory) 

T. Himasthla harrisoni S.J. Johnston, 1917 
Qld: SJJ, 1917, p.195 
syn. Echinostoma sp. 

Qld: THJ, 1912b, p.107 
syn. Echinostoma (Acanthochasmus) sp. 
Qld: THJ, 1916, p.46; THJ, 1918, p.211 
C. Unidentified 

Qld, NSW, SA: AHC 

A. Arhythmorhynchus johnstoni Golvan, 1960 

syn. Arhythmorhynchus frassoni of THJ & SJE, 
1951, not Molin, 1858 
Qld: THJ & SJE, 1951, p.3 
syn. Echinorhynchus sp. 

Qld: THJ, 1912b, p.107; 1914a, p.110 
Arhythmorhynchus sp. 

146. Tringa glareola (Linne) 

Wood Sandpiper (migratory) 

3/3 : CI N3 

T. Cyclocoelidae, unidentified 

C. Unidentified 

N. Tetrameres scotopacidis Mawson, 1968b 

SA: PMM, 1968b, p.381 


147. Tringa brevipes (Vieillot) 

Grey-tailed Tattler (migratory) 

1/1, N 

N. Skrjabinoclava sp. 

T. Haematotrephus sp. 

C. Unidentified 

N. Unidentified 

SA: believed lost 

150. Tringa stagnatilis (Bechstein) 

Marsh Sandpiper (migratory) 

2/2 : C2 Nl 

C. Paruierina rauschi Freeman, 1957 

N. Unidentified 

SA: AHC (immature) 

151. Tringa terek (Latham) 
Terek Sandpiper (migratory) 

1/1 : N A 

N. Skrjabinoclava decorata (Solonitzin, 1928) 

Schisrorophus cornutus Sobolev, 1943 

Viktorocara limosae Mawson, 1968 b 

Viktorocara schejkini Gushanskaya, 1950 

A. Unidentified 

SA: specimen not kept 

152. Gallinago hardwickii (Gray) 

Latham's Snipe (migratory) 
2/2 : C 

C. Haploparaxis australis (T.H. Johnston, 1913) 

syn. Aptoparaksis australis T.H. Johnston, 1913 
Qld: THJ, 1913, p.90 

SA: Cleland, 1922, p.105 
Qld: AHC 



153. Gallinago megala Swinhoe 
Swinhoe's Snipe (migratory) 

C. Haploparaxis australis (T.H. Johnston, 1913) 
Qld: AHC 

154. Limosa lapponica (Linne! 
Bar-tailed Godwit (migratory) 

l/l : C N 

T. Cyclocoelum taxorchis S.J. Johnston, 1917 

NSW: SJJ, 1917, p.239 
Cvclocoetum sp. 

Acanthoparvphium marilae Yamaguti, 1934 

Qld: Bearup, I960, p.219 
Acanthoparvphium squatarolae Yamaguti, 1934 

Qld: Bearup, 1960, p.219 
Himasthla ? megacotyia Yamaguti, 1939a 

Platynotrema sp. 

Maritrema sp. 

Heterophyidae, unidentified 

C. Unidentified 


N. Srreptocara sp. 

Schistorophus limosae Mawson, 1968b 

Qld: Mawson, 1968b, p.287 
Viktorocara limosae Mawson, 1968b 

Qid: PMM, 1968b, p.287 
A. A rhythmorhynchus limosae Edmonds, 1971 
'Qld: Edmonds, 1971, p.58 


155. Calidris canutus (Linne) 
Red Knot (migratory) 

8/7 : T7 N7 

T. Cycloeoelidae, unidentified 

Acanthoparvphium marilae Yamaguti, 1934 

Qld: Bearup, I960, p.219 
Cloeophora sp. [spines ? (21)] 

Himasthlinae [spines ? (31(4))] 

N. Tetrameres calidris Mawson, 1968b 

NT: PMM, 1968b, p.30() 
Sciadiocura umbellifera (Molin, 1860) 

NT: PMM, 1968b, p.285 
Schistorophus longicornis (Hemprich & Ehrenberg, 
in Schneider, 1866) 

NT: PMM, 1968b, p.285 
Viktorocara sp. 

NT: PMM, 1968b, p.289 

156. Calidris lenuirostris (Horsfield) 

Great Knot (migratory) 

T. Acanthoparvphium marilae Yamaguti, 1934 

Qld: Bearup, 1960, p.219 
Acanthoparvphium squatarolae Yamaguti, 1934 

Qld: Bearup, 1960, p.219 
Himasthla kusasigi Yamaguti, 1939a 


Qld: CIHUS (? from pancreas) 

157. Calidris acuminata (Horsfield) 
Sharp-tailed Sandpiper (migratory) 

32/15 : T4 C12 N7 

T. ? Notocotylidae, unidentified 

SA: AHC (immature) 
Echinostomatinae [spines (45(5))| 

SA: AHC (immature) 
Echinostomatidae [spines ? (37)] 

SA: AHC (immature) 
Endocotvle incana Belopolskava, 1952 

Qld: Deblock & Pearson," 1968b, p.463 
? Microphallinae, unidentified 

SA: believed lost 

SA: AHC (immature) 

SA: AHC (cysts) 
C. Trichocephaloides sp. 


NSW: Cleland, 1922, p. 105 (AHC) 

N. Tetrameres scolopacidis Mawson, 1968b 

SA: PMM, 1968b, p.30l 

158. Calidris ruficollis (Pallas) 

Red-necked Stint (migratory) 

29/28 : C23 N15 Al 

T. Microphallidae, unidentified 
Tas: AHC 
? Heterophvidae, unidentified 

Ta>: AHC 
Trematoda^ unidentified 
Tas: AHC (cyst) 
C. Unidentified 

Tas, SA: AHC 
N. Stellocaronema glareolae Mawson, 1968b 
Tetrameres scolopacidis Mawson, 1968b 
SA: PMM, 1968b, p.301 
Tas: AHC 
Tetrameres sp. 
Tas: AHC 
Echinuriu heterobrachiata Wehr, 1937 

SA: PMM, 1968b, p.289 
Skrjabinoclava sp. 
A. Unidentified 

Tas: AHC 


REC, S. AUST. MUS. 19(15): 219-325 

August, I9S6 

159. Calidris ferruginea (Pontoppidan) 

Curlew Sandpiper (migratory) 

3/1 : C 

C. Unidentified 

160. Caiidris alba (Pallas) 

Sanderling (migratory) 

l/l : C 

C. Unidentified 

161. Limicoia falcinellus (Pontoppidan) 

Broad-billed Sandpiper (migratory) 


No helminths recorded 


162. Stiltia Isabella V id Hot 

Australian Pratincole (migratory) 
2/2 : C2 Nl 

C. Unidentified 

N. Stellocaronema glareolae Mawson, 1968b 

SA: PMM, 1968b, p.296 


163. Larus novaehollandiae Stephens 

Silver Guli 

36/36 : T22 C27 N8 A9 

T. Austrohilharzict terrigafensis S.J. Johnston, 1917 

Qld: Rohde, 1977, p.39 

NSW: SJJ, 1917, p.235 

NSW: Bearup, 1956, p.471 

SA: THJ, 1941, p.282 

WA: Appleton, 1983, p.249 

Vic, SA: AHC 
Gigantobtlharzia sp. 

Qld: Rohde, 1978, p.40 
Schistosomal idae, unidentified 

Cloacitrema narrabeenense Howell & Bearup, 1967 

NSW: Howell & Bearup, 1967, p.182 
Parorchis acanthus (Nicoll, 1906) 

SA: LMA, 1954, p.173 

Philophthalmus burrili Howell & Bearup, 1967 

NSW: Howell & Bearup, 1967, p.182 
Acanthoparyphium spinidosum SJ. Johnston, 1917 

NSW: Bearup, 1960, p.219 (exptl) 
? Echinoparyphium sp. [spines (35(5))] 

Echinostoma sp. [spines missing] 

Stephanoprora sp. 

Tas: AHC 

Fxhinostomatinae [spines (33(5))] 

Fchinostomatinae [spines (37(4-5))] 

Prosthogonimus sp. 

Gynoecotyla brisbanensis Deblock & Pearson, 

Qld: Deblock & Pearson, 1968a, p.141 
Levinseniella sp. 

Microphallus papillornatus Deblock & Pearson, 

Qld: Deblock & Pearson, 1969, p.396 
Pachytrema calculus Looss, 1907 

NSW, SA: LMA, 1971, p.105 

Vic: AHC 
Galactosomum angelae Pearson, 1973 

SA: Pearson, 1973, p.364 
Galactosomum bearupi Pearson, 1973 

Qld: Pearson, 1973, p.370 
Galactosomum ussuriense Oshmarin, 1963 

Qld: Pearson, 1973, p.429 
Heterotestophves sp. 

Stictodora diplacaniha T.H. Johnston, 1942a 

Stictodora lari Yamaguti, 1939a 

NSW: Bearup, 1961, p.251 (exptl & natural) 
syn. Stictodora sp. 
" NSW: Bearup, 1958, p.219 (exptl) 
Diplostomum spathaceum (Rudolphi, 1819) 
syn. Diplostomum murrayense (T.H. Johnston & 
Cleiand. 1938) 

SA: THJ & LMA, 1941a, p.140 

SA: Dubois & LMA, 1972, p.208 
Diplostomum sp. 

Neodiplostomum pricei Krull, 1934 

Washington DC, USA (cage): Krull, 1934, 
Cardiocephaloides hil/ii (SJ. Johnston, 1904) 

SA: Dubois & LMA, 1972, p.201 

Tas: AHC 
syn. Uolostomum hillii S.J. Johnston, 1904 

NSW: SJJ, 1904, p.lll 
C. Tetrahothrius sp. 

Anomotaenia hydrochelidonis Dubinina, 1^54 


NSW, Tas, SA: AHC 

Vic: UMVS 
N. Capillaria jaenschi T.H. Johnston & Mawson, 

SA: THJ & PMM, 1945b, p.245 
Capillaria laricola Wassiiikova & Gushanskava, 

SA: THJ & PMM, 1947, p.549 
Capillaria thomascameroni Mawson, 1969 

SA: PMM, 1969, p.1104 
Tetrameres sp. 

Tas, SA: AHC 



Pectinospirura argentata Wehr, 1933 

Tas: AHC 
Eufilaria sp. 

SA: PMM, 1969, p.1110 
Filarial worm, unidentifiable 

Tas: AHC 
Microfilaria sp. 

London Zoo: Scott, 1926, p.237 
A. FilicoUis sphaerocephalus (Bremscr, 1819) 

SA: THJ & SJE, 1947, p.255 

164. Chlidonias hybrida (Pallas) 

Whiskered Tern 

20/18 : T13 C7 N8 

T. ? Ornithobilharzia sp. 
Echinostomatidae, [spines (41-43(5))] 

SA: AHC (metacercariae) 
Prosthogonimus vitellatus Nicoll, 1914a 

SA: LMA, 1973, p.859 
Pachvtrema calculus Looss, 1907 

SA: LMA, 1971, p.105 
Diplostomum spathaceum (Rudolphi, 1819) 
syn. Diplostomum murrayense (T.H. Johnston & 
Cleland, 1938) 
SA: THJ & LMA, 1941a, p.140 
Cardiocephaloides musculosus (S.J. Johnston, 
SA: Dubois & LMA, 1972, p.202 

C. Unidentified 

N. Capillaha jaenschi T.H. Johnston & Mawson, 
SA: THJ & PMM, 1945b, p.245 
Contracaecum sp. 

SA: THJ & PMM, 1947, p.551 
Tetrameres sp. 
Chevreuxia aust rails T.H. Johnston & Mawson, 
SA: THJ & PMM, 1941c, p.259 
Cosmocephalus jaenschi T.H. Johnston & Mawson, 
Acuaria fs.l.) sp. 

SA: THJ & PMM, 1941e, p.260 
Streptocara recta (Linstow, 1879) 

SA: THJ & PMM, 1942e, p.71 
Spirurida, unidentified 
SA: AHC (larva) 

165. Gelochelidon miotic j (Gmelin) 
Gull-billed Tern 

1/1 : N 

N. Habronematinae, unidentified 
SA: PMM, 1969, p.1110 
Tetrameres sp. 

SA: PMM, 1969, p.1109 







166. Hydroprogne caspia (Pallas) 

Caspian Tern (migratory) 

6/6 : Tl C3 N5 Al 

Echinostoma sp. 

Echinostomatinae [spines (45(5))] 

SA: AHC (immature) 
Galactosomum angelae Pearson, 1973 

SA: Pearson, 1973, p.364 
Galactosomum bearupi Pearson, 1973 

Qld: Pearson, 1973, p.370 
Galactosomum ussunense Oshmarin, 1963 

Qld: Pearson, 1973, p.429 
Galactosomum sp. 


SA: AHC (2 spp.) 
Heterophyidae, unidentified 

Diplostomum parvulum Dubois & Angel, 1972 

SA: Dubois & LMA, 1972, p.206 
Posthodiplostomum australe Dubois, 1937a 

SA: Dubois & LMA, 1972, p.212 
Cardiocephaloides musculosus (S.J. Johnston, 

Qld: Dubois & LMA, 1972, p.202 


Trichostrongylus incertus T.H. Johnston & 
Mawson, 1941e 

SA: THJ & PMM, 1941c, p.254 
Streptocara pectinifera Neumann, 1900 

Acuariidae, unidentified 

SA: THJ & PMM, 1941e, p.260 (larva) 


SA: AHC (poor specimen) 

167. Sterna sumatrana Raffles 

Black-naped Tern 


No helminths recorded 

168. Sterna fuscata Linne 
Sooty Tern 

2/1 : N 

Galactosomum bearupi Pearson, 1973 

Qld: Pearson, 1973, p.370 
Galactosomum renincolum Pearson, 1973 

Qld: Pearson, 1973, p.412 
Galactosomum ussuriense Oshmarin 1963 

Qld: Pearson, 1973, p.429 
Seuratia shipleyi (Stossich, 1900) 

Qld, NSW: AHC 


REC S. AUST. MUS. 19(15): 219-325 

August, 19S6 

169. Sterna albifrons Pallas 
Little Tern 

T. Stictodora sp. cf. S. manilensis Africa & Garcia, 
Qld: Pearson, 1960, p.93 

170. Sterna nereis (Gould) 

Fairy Tern 

2/1 : A 

A. Unidentified 

171. Sterna hergii Lichtenstein 

Crested Tern 
5/4 : T3 C3 N3 

T, Austrobilharzia terrigatensis (S.J. Johnston, 1917) 
NSW: Bearup, 1955, p.955 
Brachylecithum megastomum (S.J. Johnston, 1917) 
syn. Lyperosomum megastomum S.J. Johnston, 
NSW: SJJ, 1917, p.225 
Gaiactosomum angelae Pearson, 1973 

SA: Pearson, 1973, p.36 
Heterophyinae, unidentified 

Qld: C1HUS 
Cardiocephaloides musculosus (S.J. Johnston, 
syn. Holostomum musculosum S.J. Johnston, 
NSW: SJJ, 1904, p. 112 
Maritrema sp. cf. eroliae Yamaguti, 1939a 
C. Tetrabothhus sp. 

Qld, NSW (cage): AHC 

SA: Cleland, 1922, p.105 (AHC) 

N. Gapillaria sp. 

Anisakis sp. 

SA: AHC (immature) 
Cosmoeephalus jaenschi T.H. Johnston & Mawson, 

Tas: AHC 
Skrjabinoeerea sp. 

Stegophorus sp. 


172. Sterna bengalensis Lesson 

Lesser Crested Tern 

T. Gaiactosomum bearupi Pearson, 1973 
Qld: Pearson, 1973, p.370 

173. Anous stolidus (Linne) 
Common Noddy 

2/1, N 

N. Contracaecum magnicollare T.H. Johnston & 
Mawson, 1941c 
Qld: THJ & PMM, 1941c, p.114 
Coral Sea: PMM, 1969, p. 11 04 
Seuratia shipieyi (Stossich, 1900) 
Coral Sea: PMM, 1969, p.1108 

174. Anous minutus Boie 

Black Noddy 

2/2 : N 

T. Gaiactosomum bearupi Pearson, 1973 
Qld: Pearson, 1973, p.371 
Gaiactosomum renincolum Pearson, 1973 
Qld: Pearson, 1973, p.412 
N. Anisakis sp. 

syn. Stomachus sp. (larvae) 
Qld: THJ & PMM, 1951, p.294 
Contracaecum magnicollare T.H. Johnston & 
Mawson, 1941c 
Qld: AHC 
Acuariidae, unidentified 
Qld: AHC (larva) 


175. Macropygia phasianella (Temminck) 
Brown Cuckoo-Dove 

C. Unidentified 

Qld: AHC 

176. Geopelia placida Gould 

Peaceful Dove 

5/1 : C 

C. Unidentified 

NT: not kept 

177. Geopelia euneata (Latham) 

Diamond Dove (endemic) 


No helminths recorded 

178. Geopelia humeralh -(Temminck) 
Bar-shouldered Dove 

4/1 : N 

N. A sea rid ia sp. 

ACT (cage): AHC (poor specimen) 

179. Chalcophaps indiea (Linne) 
Emerald Dove 

C. Unidentified 

Qld: Cleland, 1922, p. 104 



180. Phaps chaleoptera (Latham) 
Common Bronzewing (endemic) 


C. Unidentified 

Vic: AHC 
N. Ascaridia columbae (Gmelin, 1790) 

Vic (cage): AHC 

181. Phaps elegans (Temminck) 

Brush Bronzewing (endemic) 

2/1 : C 

C. Unidentified 

SA: Cleland, 1922, p. 104 

182. Ocyphaps lophotes (Temminck & Laugier) 

Crested Pigeon (endemic) 

20/2 : C 

C. Unidentified 

183. Petrophassa albipennis Gould 

While-quilled Rock-pigeon (endemic) 

No helminths recorded 

184. Geophaps ptumifera Gould 

Spinifex Pigeon (endemic) 


N. Microfilaria sp. 

London (cage): Plimmer, 1912, p.408 

185. Geophaps seripta (Temminck) 

Squatter Pigeon (endemic) 


C. Unidentified 

Qld: AHC 

186. Geophaps smithii (Jardinc & Selby) 

Partridge Pigeon (endemic) 

C. Unidentified 

187. Leueosareia melanoleuca (Latham) 

Wonga Pigeon (endemic) 

T. Brachvlaima pu/chellum (S.J. Johnston, 1917) 
Qld: AHC 
syn. Harmosiomum pulchellum S.J. Johnston, 
NSW: SJJ, 1917, p.227 
Brachvlaima sp. 
? loc: AHC 
C. Davainea sp. 

NSW: THJ, 1912b, p.106 
Qld: AHC 

Qld: AHC 
N. Odontoterakis bancrofti (T.H. Johnston, 1912a) 
syn. Heterakis bancrofti T.H. Johnston, 1912a 
Qld: THJ Sl PMM," 1941d, p.251 


188. Calyptorhynvhus magnificus (Shaw) 

Red-tailed Black-Cockatoo (endemic) 

1/1 : C 

C. Unidentified 

NT: specimens not kept 

189. Calyptorhynchus funereus (Shaw) 
Yellow-tailed Black-Cockatoo (endemic) 


N. Syngamus trachea (Montagu, 1811) 

Vic (cage): Harrigan & Arundel, 1978, p.360 

190. Callocephalon fimbriatum (Grant) 

Gang-gang Cockatoo (endemic) 

N. Ascaridia platyceri Hartwich & Tscherner, 1979 
SA (cage): AHC 

191. Cacatua roseicapilla Vieillot 

Galah (endemic) 


C. Hemiparonia sp. 

SA (? cage): AHC 
Raillietina leptosoma (Diesing, 1850) 
syn. Davainea leptosoma Diesing, 1850 
Aust.: Fuhrmann, 1908, p.160 
N. Ascaridia sp. 

New Zealand: Weekes, 1982, p.457 

192. Caeatua tenuirostris (Kuhl) 

Long-billed Corella (endemic) 


No helminths recorded 

193. Caeatua sp. 

A corella 

N. Aprocta sp. 

Qld: AHC 

194. Caeatua sanguinea Gould 

Little Corella 


N. Microfilaria sp. 

Qld: Mackerras, 1962, p.436 


REC. S. AUST. MUS. 19(15): 219-325 

August, 1986 

195. Cacatua leadbeateri (Vigors) 
Pink Cockatoo (endemic) 


C. Hemiparonia cacatuae (Maplestone, 1922b) 

syn. Hemiparonia merotomochaeta Woodland, 
Aust: Woodland, 1930, p.221 
N. Heterakis gallinarum (Schrank, 1788) 

syn. Heterakis gallinae (Gmelin, 1790) 
NSW (cage): THJ & PMM, 1941d, p.251 

196. Cacatua galerita (Latham) 

Sulphur-crested Cockatoo 


T. Platynosomum proxillicens (Canavan, 1937) 

Fla, Wis, USA (cage): Kazakos et al, 1980, 
syn. Dicrocoelium proxillicens Canavan, 1937 

Penn, USA (cage): Canavan, 1937, p.478 
syn. Trematodes' 

Penn, USA (cage): Ratcliffe, 1933, p.21 
syn. Platynosomum ventroplicatum Heidegger & 

Mendheim, 1938b 
syn. P. fallax Heidegger & Mendheim, 1938a 
Germany (cage): Heidegger & Mendheim, 
1938a, p.94; 1938b, p.674 
Dicrocoeliinae, unidentified 
Cal. USA (cage): AHC 
syn. Dicrocoeliidae, unidentified 

Cal. USA (cage): Koch & Duhamel, 1982, 
C. Hemiparonia cacatuae (Maplestone, 1922b) 
Qld: Schmidt, 1972, p.1092 
SA (cage): AHC 
syn. Schizotaenia cacatuae Maplestone, 1922b 
Qld: Maplestone, 1922b, p.305 
Raillietina cacatuina (T.H. Johnston, 1913) 
Qld: Schmidt, 1972, p.1088 
Indonesia (cage): Sukarish, 1981, p.53 
syn. Davainea cacatuina T.H. Johnson, 1913 
Qld: THJ 1913, p.79 
Raillietina paucitesticulata (Fuhrmann, 1908) 

Aust: BM(NH) 

NSW (cage): Cleland, 1922, p.105 (AHC) 
Qld: AHC 
N. Microfilaria sp. 

Qld: Mackerras, 1962, p.436 

198. Geoffroyus geoffroyi (Bechstein) 
Red-cheeked Parrot 

N. Cardiofilaha dubia (T.H. Johnston & Mawson, 
syn. Carinema dubia T.H. Johnston & Mawson, 
Qld: THJ & PMM, 1940a, p.357 
syn. ^Filarial worm' 

Qld: Cleland, 1922, p. 107 (AHC) 


199. Triehoglossus haematodus (Linne) 

Rainbow Lorikeet 


C. Paronia trichoglossi (Linstow, 1888) 
Qld: Schmidt, 1972, p.1092 

Tas: Fuhrmann, 1908, p.159; 1932, p.283 (no 
detail given, may be repetition of Linstow's 
record, with mistaken locality for 'Cape 
syn. Moniezia trichoglossi (Linstow, 1888) 

Qld: THJ, 1913, p.78 
syn. Taenia trichoglossi Linstow, 1888 
Qld: Linstow, 1888, p.14 

Qld: Cleland, 1922, p.105 
N. Microfilaria sp. 

Qld: Bancroft, 1889, p.61 

200. Glossopsitta concinna (Shaw) 

Musk Lorikeet (endemic) 


N. Microfilaria sp. 

NSW: Mackerras, 1962, p.436 

201. Glossopsitta porphyrocephala (Dietrichsen) 

Purple-crowned Lorikeet (endemic) 


No helminths recorded 

202. Glossopsitta pusilla (White) 

Little Lorikeet (endemic) 

N. Microfilaria sp. 

Qld: Cleland & THJ, 1912, p.430 


197. Eclectus roratus (P.L.S. Muller) 
Eclectus Parrot 

C. Unidentified 

Qld: AHC 
N. Ascaridia columbae (Gmelin, 1790) 

SA (cage): PMM, 1985, p.191 


203. Alisterus scapularis (Lichtenstein) 
Australian King Parrot (endemic) 

N. Ascaridia columbae (Gmelin, 1790) 

Qld (cage): Mines & Green, 1983, p.279 
Ascaridia platyceri Hartwich & Tscherner, 1979 
Qld (cage), Vic (cage), SA (cage): PMM, 1985, 



204. Aprosmictus erythropterus Gmelirt 
Red-winged Parrot 

N. Ascuridia ptatyceri Hartwich & Tscherner, 1979 
syn. Ascuridia sprenti Mines, 1979 
" Vic (cage): Mines, 1979, p.374 

205, Polytelis swainsonii (Desmarest) 
Superb Parrot (endemic) 


N. Ascuridiu ptutyceri Hartwich & Tscherner, 1979 
Tas (cage): PMM, 1985, p.l9l 

206. Polytelis anthopephis (Lear) 
Regent Parrot (endemic) 

C. Cotugnia polylelidis Burt, 1940 

Ceylon (cage); Burt, 1940, p.65 
N. Ascuridia ptatyceri Hartwich & Tscherner, 1979 
Berlin (cage): Hartwich & Tscherner, 1979, p.64 
syn. Ascuridia sprenti Mines* 1979 
Vic (cage): Mines, 1979, p.374 
Ascuridia sp. 

New Zealand (cage): Weekes, 1983, p.457 

207. Polytelis alexandrae Gould 
Princess Parrot (endemic) 

N. Ascuridia columbae (Gmelin, 1790) 

Old (cage): Mines & Green, 1983, p.279 
NSW (cage): THJ & PMM, !941d, p.253 
Ascaridia platvceri Hartwich 8c Tscherner, 1979 
Vic (cage), SA (cage): PMM, 1985, p.191 
syn. Ascuridia sprenti Mines, 1979 
' Vic (eage): Mines, 1979, p.371 

208. Nymphicus hotlandicus (Kerr) 

Cockatiel (endemic) 

N. Hluriu (sj.) sp. 

NSW: THJ & PMM, 1940a, p.361 
Buylisuscaris sp. 

Iowa, USA (cage): Myers, Monroe & Greeve, 
1983, p.1089. 
Ascaridia platvceri Hartwich & Tscherner, 1979 

Berlin (cage): Hartwich & Tscherner, 1979, p.64 

SA (cage): AHC 
Ascaridia sp. 

New Zealand (cage): Weekes, 1983, p.457. 


209. Mclopsittacus undulatus (Shaw) 

Budgerigah (endemic) 


T. Austrobilharzia terrigalensis SJ. Johnston, 1917 
NSW: Bearup, 1956, p.471 (exptl.) 
Trichobilhurziu sp. 

NSW: CTHUS (exptl) 

N. Cupillaha sp. 

Qld (? cage): AHC 
Ascuridia columbae (Gmelin, 1790) 

Qld (cage); Mines & Green, 1983, p.279 
Ascaridia platvceri Hartwich & Tscherner, 1979 

Vic (cage), SA (cage): PMM, 1985, p.191 
Ascaridia sp. 

New Zealand (cage); Weekes, 1983, p.457 
Procymeu incerta (A.J. Smith, 1908) 
syn. Spiropferu incerta A.J. Smith, 1908 

USA (cage): A.J. Smith, 1908, p.269 

210. Lathamus discolor (White) 

Swift Parrot (endemic) 

N. Ascaridia columbae (Gmelin, 1790) 
SA (cage): PMM, 1985, p. 191 
Ascaridia platvceri Hartwich & Tscherner, 1979 
SA (cage): PMM, 1985, p.191 

211. Platycereus caledonicus (Gmelin) 
Green Rosella (endemic) 

3/1 : N 

N. Unidentified 

Tas: AHC (larva) 

212. Platycercus elegans (Gmelin) 

Crimson Rosella (endemic) 

29/4 : C3 Nl 

C. Hemiparoniu bancrofti (TH. Johnston, 1912c)